Endangered and Threatened Wildlife and Plants; Endangered Status for Physaria globosa (Short's bladderpod), Helianthus verticillatus (whorled sunflower), and Leavenworthia crassa (fleshy-fruit gladecress), 47109-47134 [2013-18213]
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Federal Register / Vol. 78, No. 149 / Friday, August 2, 2013 / Proposed Rules
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS–R4–ES–2013–0087;
4500030113]
RIN 1018–AZ11
Endangered and Threatened Wildlife
and Plants; Endangered Status for
Physaria globosa (Short’s bladderpod),
Helianthus verticillatus (whorled
sunflower), and Leavenworthia crassa
(fleshy-fruit gladecress)
Fish and Wildlife Service,
Interior.
ACTION: Proposed rule.
AGENCY:
We, the U.S. Fish and
Wildlife Service, propose to list
Physaria globosa (Short’s bladderpod),
Helianthus verticillatus (whorled
sunflower), and Leavenworthia crassa
(fleshy-fruit gladecress) as endangered
under the Endangered Species Act of
1973, as amended (Act). If we finalize
this rule as proposed, it would extend
the Act’s protections to Physaria
globosa (Short’s bladderpod),
Helianthus verticillatus (whorled
sunflower), and Leavenworthia crassa
(fleshy-fruit gladecress) to conserve
these species.
DATES: We will accept all comments
received or postmarked on or before
October 1, 2013. Comments submitted
electronically using the Federal
eRulemaking Portal (see ADDRESSES
section, below) must be received by
11:59 p.m. Eastern Time on the closing
date. We must receive requests for
public hearings, in writing, at the
address shown in the FOR FURTHER
INFORMATION CONTACT section by
September 16, 2013.
ADDRESSES: You may submit comments
by one of the following methods:
(1) Electronically: Go to the Federal
eRulemaking Portal: https://
www.regulations.gov. In the Search
field, enter Docket No. FWS–R4–ES–
2013–0087, which is the docket number
for this rulemaking. Then, in the Search
panel on the left side of the screen,
under the Document Type heading,
click on the Proposed Rules link to
locate this document. You may submit
a comment by clicking on ‘‘Comment
Now!’’ If your comments will fit in the
provided comment box, please use this
feature of https://www.regulations.gov, as
it is most compatible with our comment
review procedures. If you attach your
comments as a separate document, our
preferred file format is Microsoft Word.
If you attach multiple comments (such
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SUMMARY:
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as form letters), our preferred format is
a spreadsheet in Microsoft Excel.
(2) By hard copy: Submit by U.S. mail
or hand-delivery to: Public Comments
Processing, Attn: FWS–R4–ES–2013–
0087; Division of Policy and Directives
Management; U.S. Fish and Wildlife
Service; 4401 N. Fairfax Drive, MS
2042–PDM; Arlington, VA 22203.
We request that you send comments
only by the methods described above.
We will post all information received on
https://www.regulations.gov. This
generally means that we will post any
personal information you provide us
(see the Information Requested section
below for more details).
FOR FURTHER INFORMATION CONTACT:
Mary E. Jennings, Field Supervisor, U.S.
Fish and Wildlife Service, Tennessee
Ecological Services Field Office, 446
Neal Street, Cookeville, TN 38501; by
telephone 931–528–6481; or by
facsimile 931–528–7075. Persons who
use a telecommunications device for the
deaf (TDD) may call the Federal
Information Relay Service (FIRS) at
800–877–8339.
SUPPLEMENTARY INFORMATION:
Executive Summary
Why we need to publish a rule. Under
the Act, if we intend to list a species are
endangered or threatened throughout all
or a significant portion of its range, we
are required to promptly publish a
proposal in the Federal Register to list
the species as endangered or threatened
and make a determination on our
proposal within 1 year. Listing a species
as an endangered or threatened species
can only be completed by issuing a rule.
This rule proposes to add three plants
to the Federal List of Endangered and
Threatened Plants. We are proposing to
list Short’s bladderpod, whorled
sunflower, and fleshy-fruit gladecress as
endangered species under the Act.
Elsewhere in today’s Federal Register,
we propose to designate critical habitat
for the Short’s bladderpod, freshy-fruit
gladecress, and the whorled sunflower.
The basis for our action. Under the
Act, we may determine that a species is
an endangered or threatened species
based on any of five factors: (A) The
present or threatened destruction,
modification, or curtailment of its
habitat or range; (B) overutilization for
commercial, recreational, scientific, or
educational purposes; (C) disease or
predation; (D) the inadequacy of
existing regulatory mechanisms; or (E)
other natural or manmade factors
affecting its continued existence.
We have determined that listing is
warranted for these species, which are
currently at risk throughout all of their
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47109
respective ranges due to threats related
to:
• For Short’s bladderpod, potential
future construction and ongoing
maintenance of transportation rights-ofway; prolonged inundation and soil
erosion due to flooding and water level
manipulation; overstory shading due to
forest succession and shading and
competition from invasive, nonnative
plant species; and small population
sizes.
• For whorled sunflower, mechanical
or chemical vegetation management for
industrial forestry, right-of-way
maintenance, or agriculture; shading
and competition resulting from
vegetation succession; limited
distribution and small population sizes.
• For fleshy-fruit gladecress, loss of
habitat due to residential and industrial
development; conversion of agricultural
sites for use as pasture; mowing and
herbicide treatment prior to seed
production; and off-road vehicles and
dumping.
We will seek peer review. We are
seeking comments from knowledgeable
individuals with scientific expertise to
review our analysis of the best available
science and application of that science
and to provide any additional
information to improve this proposed
rule. Because we will consider all
comments and information we receive
during the comment period, our final
determinations may differ from this
proposal.
Information Requested
We intend that any final action
resulting from this proposed rule will be
based on the best scientific and
commercial data available and be as
accurate and as effective as possible.
Therefore, we request comments or
information from other concerned
governmental agencies, Native
American tribes, the scientific
community, industry, or any other
interested parties concerning this
proposed rule. We particularly seek
comments concerning:
(1) The species’ biology, range, and
population trends, including:
(a) Habitat requirements for feeding,
reproducing, and sheltering;
(b) Genetics and taxonomy;
(c) Historical and current range,
including distribution patterns;
(d) Historical and current population
levels, and current and projected trends;
and
(e) Past and ongoing conservation
measures for these species, their
habitats or both.
(2) The factors that are the basis for
making a listing determination for a
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species under section 4(a) of the Act,
which are:
(a) The present or threatened
destruction, modification, or
curtailment of its habitat or range;
(b) Overutilization for commercial,
recreational, scientific, or educational
purposes;
(c) Disease or predation;
(d) The inadequacy of existing
regulatory mechanisms; or
(e) Other natural or manmade factors
affecting its continued existence.
(3) Biological, commercial trade, or
other relevant data concerning any
threats (or lack thereof) to this species
and regulations that may be addressing
those threats.
(4) Additional information concerning
the historical and current status, range,
distribution, and population size of
these species, including the locations of
any additional populations of these
species.
(5) Current or planned activities in the
areas occupied by these species and
possible impacts of these activities on
them.
Please note that submissions merely
stating support for or opposition to the
action under consideration without
providing supporting information,
although noted, will not be considered
in making a determination, as section
4(b)(1)(A) of the Act directs that
determinations as to whether any
species is an endangered or threatened
species must be made ‘‘solely on the
basis of the best scientific and
commercial data available.’’
You may submit your comments and
materials concerning this proposed rule
by one of the methods listed in the
ADDRESSES section. We request that you
send comments only by the methods
described in the ADDRESSES section.
If you submit information via https://
www.regulations.gov, your entire
submission—including any personal
identifying information—will be posted
on the Web site. If your submission is
made via a hardcopy that includes
personal identifying information, you
may request at the top of your document
that we withhold this information from
public review. However, we cannot
guarantee that we will be able to do so.
We will post all hardcopy submissions
on https://www.regulations.gov. Please
include sufficient information with your
comments to allow us to verify any
scientific or commercial information
you include.
Comments and materials we receive,
as well as supporting documentation we
used in preparing this proposed rule,
will be available for public inspection
on https://www.regulations.gov, or by
appointment, during normal business
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hours, at the U.S. Fish and Wildlife
Service, Tennessee Ecological Services
Field Office (see FOR FURTHER
INFORMATION CONTACT).
Background
Previous Federal Actions
The Act requires the Service to
identify species of wildlife and plants
that are endangered or threatened, based
on the best available scientific and
commercial data. The Act directed the
Secretary of the Smithsonian Institution
to prepare a report on endangered and
threatened plant species, which was
published as House Document No.
94–51. The Service published a notice
in the Federal Register on July 1, 1975
(40 FR 27824), in which we announced
that more than 3,000 native plant taxa
named in the Smithsonian’s report and
other taxa added by the 1975 notice
would be reviewed for possible
inclusion in the List of Endangered and
Threatened Plants. The 1975 notice was
superseded on December 15, 1980 (45
FR 82480), by a new comprehensive
notice of review for native plants that
took into account the earlier
Smithsonian report and other
accumulated information. On November
28, 1983 (48 FR 53640), a supplemental
plant notice of review noted the status
of various taxa. Complete updates of the
plant notice were published on
September 27, 1985 (50 FR 39526) and
on February 21, 1990 (55 FR 6184).
In these reviews, Short’s bladderpod
(as Lesquerella globosa) was listed as a
Category 2 candidate, taxa for which
information in the possession of the
Service indicated that proposing to list
the species as endangered or threatened
was possibly appropriate, but for which
sufficient data on biological
vulnerability and threat were not
available to support listing rules.
Further biological research and field
study usually was necessary to ascertain
the status of taxa in this category.
Fleshy-fruit gladecress was
recognized as consisting of two varietal
taxa in these reviews, Leavenworthia
crassa var. crassa and L. crassa var.
elongata. In the 1980 review, var. crassa
was listed as a Category 2 candidate,
while var. elongata was listed as a
Category 1 candidate, taxa for which the
Service had sufficient information to
support listing as either endangered or
threatened. In the 1983, 1985, and 1990
reviews both varieties of Leavenworthia
crassa were listed as Category 2
candidates. Many Category 2 candidate
species were found not to warrant
listing, either because they were not
endangered or threatened or because
they did not qualify as species under the
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definitions in the Act (58 FR 51144,
September 30, 1993).
In 1993, the Service eliminated
candidate categories, and Short’s
bladderpod and the two varieties of
fleshy-fruit gladecress were no longer
candidates until they were again
elevated to candidate status on October
25, 1999 (64 FR 57534). The 1999
review elevated the species
Leavenworthia crassa (fleshy-fruit
gladecress) to candidate status, but did
not recognize intraspecific taxa
(varieties) due to changes in
scientifically accepted taxonomy.
Whorled sunflower was first listed as a
candidate species in the 1999 review.
All three of these species were then
included in subsequent candidate
notices of review on October 30, 2001
(66 FR 54808), June 13, 2002 (67 FR
40657), May 4, 2004 (69 FR 24876), May
11, 2005 (70 FR 24870), September 12,
2006 (71 FR 53756), December 6, 2007
(72 FR 69034), December 10, 2008 (73
FR 75176), November 9, 2009 (74 FR
57804), November 10, 2010 (75 FR
69222), October 26, 2011 (76 FR 66370),
and November 21, 2012 (77 FR 69994).
Species Information
Short’s bladderpod
Physaria globosa is a member of the
mustard family (Brassicaceae) known
from Posey County, Indiana; Clark,
Franklin and Woodford Counties,
Kentucky; and Cheatham, Davidson,
Dickson, Jackson, Montgomery, Smith,
and Trousdale Counties, Tennessee. The
following description is based on Flora
of North America (https://
www.efloras.org/florataxon.aspx?flora_
id=1&taxon_id=250095135, accessed on
December 7, 2012) and Gleason and
Chronquist (1991, p. 187).
Short’s bladderpod is an upright
biennial or perennial (lives for 2 years
or longer) with several stems, some
branched at the base, reaching heights
up to 50 centimeters (cm) (20 inches
(in.)), and which are leafy to the base of
the inflorescence (a group or cluster of
flowers arranged on a stem that is
composed of a main branch or a
complicated arrangement of branches).
The basal leaves, borne on short petioles
(stalks) are 2.5 to 5 cm (1 to 2 in.) in
length and 0.5 to 1.5 cm (0.2 to 0.6 in.)
wide, obovate (egg-shaped and flat, with
the narrow end attached to the stalk) or
oblanceolate (with the widest portion of
the leaf blade beyond the middle) in
shape, with a smooth or slightly wavy
margin, and gray-green in color due to
a layer of dense hairs. Leaves are
gradually reduced in size and petiole
length higher up the stem. Numerous
flowers are borne on a raceme (elongate,
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spike-shaped inflorescence to which
individual flowers are attached by
slender pedicels, or stalks, which in
Short’s bladderpod are longer than the
flowers). The yellow flowers are
composed of four spoon-shaped petals,
0.4 to 0.7 cm (0.16 to 0.28 in.) long. The
fruit is globose in shape and lightly
beset with stellate (star-shaped) hairs,
but becoming smooth with time.
Taxonomy. A member of the mustard
family (Brassicaceae), Short’s
bladderpod was first described as
Vesicaria globosa by Desvaux in 1814
(Payson 1922, pp. 103–236). Because of
several distinctive characters, Watson
(1888, pp. 249–255) proposed that the
American species of the genus Vesicaria
be placed in the genus Lesquerella. This
treatment was recognized as valid, until
Al-Shehbaz and O’Kane (2002, entire)
reunited most of the genus Lesquerella
with the genus Physaria. This
determination was supported by
molecular, morphological, cytological,
biogeographic, and ecological lines of
evidence (Al-Shehbaz and O’Kane 2002,
p. 320). Flora of North America
recognizes this change, using the
scientific name Physaria globosa for
Short’s bladderpod (https://
www.efloras.org/florataxon.aspx?flora_
id=1&taxon_id=250095135, accessed on
April 20, 2011).
Distribution and Status. In a 1992
status survey for Short’s bladderpod,
Shea (1993, pp. 6–15) observed the
species at only 26 of 50 historical sites:
1 in Indiana, 14 in Kentucky, and 11 in
Tennessee. The remaining sites were
classified as follows (Shea 1993, p. 10–
14):
• Status uncertain—4 occurrences
where the species had been observed
during the prior 25 years and where
appropriate habitat existed with no
evidence that the occurrence had been
destroyed (Shea population numbers 27
through 30).
• Extirpated—one occurrence where
the habitat had been severely altered
(Shea population number 31).
• Historical—5 occurrences where the
species had not been observed during
the prior 25 years, but where
appropriate habitat remained (Shea
population numbers 32 through 36).
• Locality information incomplete—
14 occurrences for which location
information was insufficient to confirm
the species’ presence or absence, despite
searches having been attempted in some
cases (Shea population numbers 37
through 50). Many of these putative
occurrences were based on herbarium
specimens dating from the late-19th to
mid-20th centuries that contained little
information about sites from which they
were collected. Except for the
populations numbered 37, 42, and 50,
Shea (1993) searched for suitable habitat
or Short’s bladderpod plants in areas
associated with these occurrences but
did not find the species.
Later surveys found Short’s bladderpod
extant at two of these sites, Tennessee
element occurrence (EO) numbers 8 and
12, which correspond to Shea’s
population numbers 34 and 29,
respectively.
We used data provided to us by
conservation agencies in the States
where the species occurs (Indiana
Natural Heritage Data Center (INHDC)
2012, Kentucky Natural Heritage
Program (KNHP) 2012, Tennessee
(Tennessee Natural Heritage Inventory
Database (TNHID) 2012) to determine
47111
the current distribution and status of
Short’s bladderpod. Difficulty in
relating the species’ distribution at the
time of Shea’s (1993, entire) status
survey to its current distribution comes
as a result of State conservation agencies
revising the mapping of some element
occurrences in these databases. In two
instances, pairs of occurrences that Shea
(1993) considered distinct have been
combined into single element
occurrences (Table 1). Conversely,
TNHID (2012) treats as two distinct
element occurrences the two locations
that Shea (1993, p. 85, 108) mapped
together as population number 23. One
of these occurrences (TN EO number 22)
was extant as of 2012 (Table 1), while
the other (TN EO number 2) is
extirpated (Table 2). Based on current
mapping, State conservation agencies
now recognize 24 element occurrences
that correspond to populations that
Shea (1993, entire) found extant in
1992. Of these 24 occurrences, 18 were
extant in 2012. Accounting for
rediscovery of the two Tennessee
occurrences that Shea (1993, pp. 10–14)
did not find during 1992, and recent
changes in element occurrence
mapping, a total of 20 occurrences that
were documented by Shea (1993, entire)
were still considered extant as of 2012
(Table 1).
The approximate range of abundance
shown in Table 1 is primarily based on
individual plants. As a result of
location, it was impossible to enumerate
individual plants. This resulted in are
two instances where TNHID surveyed
these populations from a boat and
reported the approximate range in
clusters.
TABLE 1—LIST OF KNOWN EXTANT SHORT’S BLADDERPOD OCCURRENCES BY STATE AND COUNTY, WITH ELEMENT OCCURRENCE (EO) NUMBERS ASSIGNED BY STATE NATURAL HERITAGE PROGRAMS (INHDC (2012), KNHP (2012),
TNHID (2012)), NUMBERS ASSIGNED TO POPULATIONS REPORTED IN SHEA (1993), AND FIRST AND LAST YEARS OF
KNOWN OBSERVATIONS
EO Number
(Shea Population Number)
County
Indiana ................................
Kentucky .............................
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State
Posey .................................
Clark ...................................
Franklin ...............................
Tennessee ..........................
Woodford ............................
Cheatham ...........................
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First
observed
Last
observed
Approximate
range of
abundance
1 (1)
1 (3)
4 (11, 12)
7 (10)
11 (13)
18 (4)
22 (9)
1941–05–06
1957
1979
1981
1983
1992
1990-Pres
2012
2009–05–21
2011–04–19
2004–05–17
2003–06–01
2012–05–09
2012–05–08
3–1000s ......
2 ..................
100–500 ......
1–100 ..........
1–52 ............
20–350 ........
2–200 ..........
23 (14)
28
1 (18)
15 (17)
1990
2005–05–06
1956–03–02
1955–04–24
2011–04–26
2010–06–02
2008–04–23
2008–04–29
60–500 ........
few ..............
100s–1000s
few–20 .........
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02AUP2
Land ownership
IDNR.
Private.
Private.
Private.
Private.
City of Frankfort.
private; Kentucky State
Nature Preserves Commission.
Private.
Private.
COE; private.
COE.
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TABLE 1—LIST OF KNOWN EXTANT SHORT’S BLADDERPOD OCCURRENCES BY STATE AND COUNTY, WITH ELEMENT OCCURRENCE (EO) NUMBERS ASSIGNED BY STATE NATURAL HERITAGE PROGRAMS (INHDC (2012), KNHP (2012),
TNHID (2012)), NUMBERS ASSIGNED TO POPULATIONS REPORTED IN SHEA (1993), AND FIRST AND LAST YEARS OF
KNOWN OBSERVATIONS—Continued
EO Number
(Shea Population Number)
20–∼1500 ....
1998–05–12
1998–05–12
1935
1971–05–16
2008–04–29
2008–04–29
2012–06–15
2012–06–15
∼50 ..............
10–25 ..........
10s–1000s ..
100s–1000s
1886–04–22
2008–05–02
∼50 ..............
2008–04–29
1998–05–08
1998–05–08
1946–04–27
2008–04–29
2008–05–06
2008–05–06
2008–05–09
∼7 clusters ...
3 clusters .....
∼50 ..............
∼50 ..............
1969–04–28
2008–05–02
20–50 ..........
28
1998–04–23
2008–04–29
∼300 ............
24
3 (25)
21 (26)
Smith ..................................
Trousdale ...........................
2012–06–15
22 (23a)
Montgomery .......................
1953–04–26
32
26
27
12 (29)
Dickson ...............................
Jackson ..............................
Approximate
range of
abundance
29
30
10 (21,22)
4 (19)
Davidson; Cheatham ..........
Davidson ............................
Last
observed
8 (34)
County
First
observed
17 (16)
State
1998–05–05
1969–05–08
1992–04–30
2008–05–06
2008–05–06
2008–05–12
∼10 ..............
40–500 ........
100–250 ......
Land ownership
Town of Ashland
City; private.
COE; private.
COE; private.
Private.
private; COE
easement.
private; COE
easement.
COE.
COE.
COE.
private; COE
easement.
private; COE
easement.
private; COE
easement.
COE.
COE; private.
COE; private.
IDNR is the Indiana Department of Natural Resources.
COE is the U.S. Army Corps of Engineers.
Pres is present.
Despite the rediscovery of the two
Tennessee occurrences and the
discovery of 10 additional occurrences
since the 1992 status survey, only 26
extant occurrences of Short’s
bladderpod are known to remain due to
the loss of 10 occurrences during the
last 20 years (Table 1). Seven of the
occurrences that Shea (1993, pp. 44–71)
observed in 1992, and three others
(Kentucky EO number 27 and Tennessee
EO numbers 23 and 25) that were seen
after 1992, have since been extirpated
(Table 2). This constitutes a loss of 27
percent of all occurrences that were
extant during 1992 or later.
TABLE 2—LIST OF EXTIRPATED SHORT’S BLADDERPOD OCCURRENCES BY STATE AND COUNTY, WITH ELEMENT OCCURRENCE (EO) NUMBERS ASSIGNED BY STATE NATURAL HERITAGE PROGRAMS (INHDC (2012), KNHP (2012), TNHID
(2012)), NUMBERS ASSIGNED TO POPULATIONS REPORTED IN SHEA (1993), AND FIRST AND LAST YEARS OF KNOWN
OBSERVATIONS
EO Number
(Shea Population Number)
County
Kentucky ............................
First observed
Last observed
* 19 (2)
12 (38)
16 (37)
* 2 (6)
* 3 (8)
5 (39)
8 (27)
14 (40)
* 20 (5)
* 21 (7)
6 (42)
13 (32)
17 (28)
+ 27
10 (43)
24 (44)
25
15 (45)
* 9 (15)
14 (33)
* 9 (20)
1963–04–27
1931
1892
1979–05
1979
1880
1981
1856
1992
1992
1942
1939
1991–Pre
1990
1903
1916
1935–pre
1923
1930
1969–04–29
1974–04–16
2005–06–09
1931–05–24
1900–05–09
1992–05–04
1994–05–12
1880–06
1981–05–03
1856–05
1992–05–19
1992–05–12
1942–05–16
1939–04–27
1991–Pre
1993–05–10
1903–05–16
1916–05–13
1935–pre
1923–05–26
1992–05–19
1969–04–29
1998–04–16
10–120
n/a
n/a
11
4
n/a
∼40
n/a
21
7
n/a
n/a
n/a
1–7
n/a
1–7
n/a
n/a
2
n/a
20–29
+ 23
State
1997–05–09
1997–05–09
∼200
Bourbon .............................
Fayette ..............................
Franklin .............................
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Jessamine .........................
Tennessee .........................
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Madison .............................
Mercer ...............................
Nelson ...............................
Powell ................................
Scott ..................................
Cheatham ..........................
Davidson ...........................
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02AUP2
Abundance
Land ownership
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private.
private; COE
easement.
private.
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TABLE 2—LIST OF EXTIRPATED SHORT’S BLADDERPOD OCCURRENCES BY STATE AND COUNTY, WITH ELEMENT OCCURRENCE (EO) NUMBERS ASSIGNED BY STATE NATURAL HERITAGE PROGRAMS (INHDC (2012), KNHP (2012), TNHID
(2012)), NUMBERS ASSIGNED TO POPULATIONS REPORTED IN SHEA (1993), AND FIRST AND LAST YEARS OF KNOWN
OBSERVATIONS—Continued
State
EO Number
(Shea Population Number)
County
First observed
Last observed
+ 25
7 (31)
2 (23b)
13 (30)
18 (35)
31
20 (24)
1998–07–24
1955–04–23
1968–05–07
1975–05–25
1967–06–01
1979–04–09
1992–05–01
1998–07–24
1955–04–23
1992–04–28
1975–05–25
1967–06–01
1979–04–09
1998–04–17
Jackson .............................
Maury ................................
Montgomery ......................
Smith .................................
Abundance
5
n/a
1
n/a
n/a
........................
30
Land ownership
COE
private.
private.
private.
private.
private.
private; COE
easement.
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* Occurrences observed by Shea (1993), but which are now considered extirpated.
+ Occurrences not documented in Shea (1993) that have been observed since 1992, but which are now considered extirpated.
COE is the U.S. Army Corps of Engineers.
Pres is present.
No records exist in State-maintained
databases for seven populations that
Shea (1993, pp. 12–13) treated as
historical or lacking sufficient locality
information to verify (population
number 41 from Kentucky, and numbers
36 and 46 through 50 from Tennessee).
Therefore, Table 1 and Table 2 do not
include entries for these Shea
population numbers. Shea (1993, p. 15)
also determined that four historical
reports for the species were erroneous:
One each from Monroe County, Indiana,
and Vinton County, Ohio; and one each
from unknown counties in Kansas and
Vermont.
There are now 8 known extant
occurrences in Kentucky, 17 in
Tennessee, and 1 in Posey County,
Indiana (Table 1). Extant occurrences in
Kentucky are distributed among Clark
(1), Franklin (6), and Woodford (1)
Counties, and in Tennessee among
Cheatham (5), Davidson (2), Dickson (1),
Jackson (2), Montgomery (3), Smith (1),
and Trousdale (2) Counties. One
Tennessee occurrence straddles the
county line between Cheatham and
Davidson Counties. There are 19
occurrences in Kentucky and 10 in
Tennessee that have either been
extirpated or for which inadequate
information exists to relocate them.
Adding the seven populations that Shea
(1993, p. 12–13) treated as either
historical or lacking complete locality
information, and which are not
represented in State-maintained
databases used to create Tables 1 and 2,
these numbers rise to 20 for Kentucky
and 16 for Tennessee. Thus, there is a
total of 62 occurrences that have been
reported for Short’s bladderpod.
However, when reporting percentages of
all known occurrences that are now or
historically were in the case of
extirpated occurrences, affected by
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various threats, we only use the 55
records that have been verified and are
currently tracked in State-maintained
databases.
There are 19 extant Short’s
bladderpod occurrences that are located
on city, State, or federal lands. The
Indiana occurrence is on lands owned
by the State of Indiana and managed by
the Indiana Department of Natural
Resources (IDNR). A portion of one
occurrence in Kentucky is located in a
State nature preserve owned and
managed by the Kentucky State Nature
Preserves Commission (KSNPC), and
another occurs in a park owned by the
City of Frankfort, where access is
limited, but no specific management is
provided for the species or its habitat.
In Tennessee, there are 15 occurrences
that are entirely or partially located on
lands owned or leased by the U.S. Army
Corps of Engineers (Corps) adjacent to
the Cumberland River. Some of these
Corps lands are wildlife management
areas (WMA) cooperatively managed by
the Tennessee Wildlife Resources
Agency (TWRA). The plants at EO
numbers 29 and 32 are located in
TWRA’s Cheatham WMA, and those at
EO numbers 24 through 27 are located
in TWRA’s Cordell Hull WMA. Part of
one occurrence in Tennessee is located
on lands owned by Ashland City.
Habitat. Short’s bladderpod typically
grows on steep, rocky, wooded slopes
and talus (sloping mass of rock
fragments below a bluff or ledge) areas.
It also occurs along tops, bases, and
ledges of bluffs. The species usually is
found in these habitats near rivers or
streams and on south- to west-facing
slopes. Most populations are closely
associated with calcareous outcrops
(Shea 1993, p. 16). The Short’s
bladderpod site in Indiana, where the
species is found in a narrow strip of
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herbaceous vegetation between a road
and forested bank of a cypress slough
(M. Homoya, Natural Heritage Program
Botanist, Indiana Department of Natural
Resources (IDNR), December 2012), is
unique among populations of the
species. The occurrence in Indiana is
within the Shawnee Hills Section of the
Interior Low Plateaus Physiographic
Province (Quarterman and Powell 1978,
pp. 30–31), on a site underlain by
undifferentiated outwash from the
Wisconsinan glaciation (Indiana
Geologic Survey 2002) as opposed to the
calcareous geology on which the species
occurs in Kentucky and Tennessee. The
soil at the Indiana site is Weinbach silt
loam, which forms in acid alluvium on
river terraces, and is nearly level with
0 to 2 percent slopes (USDA 1979, p.
89). This site is on a terrace adjacent to
an oxbow swamp formed in an
abandoned meander of the Wabash
River (Quarterman and Powell 1978, p.
244).
Kentucky occurrences are located on
bluffs and hillsides adjacent to the
Kentucky River or its tributaries within
the Bluegrass Section of the Interior
Low Plateaus Province (Fenneman 1938,
pp. 411–448; Quarterman and Powell
1978, pp. 30–31). Extant occurrences in
Kentucky predominantly are found on
the Ordovician age Lexington Limestone
and Tanglewood Limestone Members
(Kentucky Geological Survey, https://
www.arcgis.com/home/item.html?id=
d32dc6edbf9245cdbac3fd7e255d3974,
accessed on January 25, 2013), and the
Fairmount-Rock outcrop Complex is the
prevalent soil type at most of the sites
where the species is found (U.S.
Department of Agriculture (USDA), Soil
Survey Geographic Database, available
online at https://soildatamart.nrcs.usda.
gov, accessed on January 30, 2013). Soils
of the Fairmount series formed from
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weathered limestone interbedded with
thin layers of calcareous shale and are
shallow, well-drained, and slowly
permeable. As implied in the name of
this complex, limestone outcrops are
common on the steeply sloped sites
where this soil occurs, especially along
river bluffs (USDA 1985, p. 64).
Tennessee occurrences are located
primarily on steep hills or bluffs
adjacent to the Cumberland River
within the Highland Rim and Central
(also known as Nashville) Basin
Sections of the Interior Low Plateaus
Province (Fenneman 1938, pp. 411–448;
Quarterman and Powell 1978, pp. 30–
31). Three occurrences in Cheatham
County are adjacent to the Harpeth
River near its confluence with the
Cumberland River. Extant occurrences
in Tennessee are found across a wider
range of geology and soils than those in
Indiana or Kentucky. The Mississippian
age Fort Payne Formation, which
includes limestone and calcareous
siltstone, and Warsaw Limestone are the
predominant geologic formations
underlying occurrences in Cheatham,
Dickson, and Montgomery Counties
(Moore et al. 1967, Wilson 1972, Marsh
et al. 1973, Finlayson et al. 1980). In
Cheatham and Dickson Counties, the
main soil mapped in locations where
Short’s bladderpod occurs is simply
‘‘Rock outcrop, very steep’’ (USDA, Soil
Survey Geographic Database, available
online at https://soildatamart.nrcs.usda.
gov, accessed on January 30, 2013). In
Montgomery County, Baxter soils and
Rock outcrop and Bodine cherty silt
loam are the soil types on which Short’s
bladderpod occurs (USDA, Soil Survey
Geographic Database, available online at
https://soildatamart.nrcs.usda.gov,
accessed on January 30, 2013). Baxter
soils formed from weathered cherty
limestone, and where they are mapped
as Baxter soils and Rock outcrop they
are steeply sloped and Rock outcrop can
make up as much as 20 percent of the
map unit (USDA 1975, pp. 12–14).
Bodine soils are well-drained, cherty
soils that formed from weathered cherty
limestone; are steeply sloped; and
include areas near the escarpment
adjacent to the Cumberland River
floodplain where cherty limestone
bedrock is exposed (USDA 1975, pp.
16–17).
Silurian age limestone and shale of
the Waynes Group and the Brassfield
Limestone and Ordovician age
limestone of the Leipers and Catheys
Formations are the predominant
geologic formations underlying the
occurrences located in Davidson County
(Wilson 1979). The dominant soils on
which Short’s bladderpod occurs in this
county are the Bodine-Sulphura
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Complex (USDA, Soil Survey
Geographic Database, available online at
https://soildatamart.nrcs.usda.gov,
accessed on January 30, 2013), which
formed from weathered cherty
limestone on sloping to very steep sites
and are somewhat excessively welldrained. Depth to bedrock within
Sulphura soils is less than 16 cm (40 in),
but deeper in Bodine soils, and chert
content is high near the surface of these
soils (USDA 1981, pp. 46–47).
Ordovician age limestones of the
Leipers and Cathey Formations, BigbyCannon Limestone, and Hermitage
Formation are the predominant geologic
formations underlying occurrences in
Smith, Trousdale, and Jackson Counties
(Wilson et al. 1972, Wilson 1975,
Wilson et al. 1980, Kerrigan and Wilson
2002). In these counties, Short’s
bladderpod occurs across a wider range
of soil series, all of which are formed
from weathered limestone or
interbedded siltstone and limestone on
steeply sloped or hilly sites. The soils
are shallow, are rocky, or contain areas
of bedrock outcrop (USDA 2001, pp. 19–
20, 28, 59, 64; USDA 2004a, pp. 22–23,
36–37, 83, 87; USDA 2004b, pp. 21, 75,
82).
Within the physical settings described
above, the most vigorous (Shea 1992, p.
24) and stable (TDEC 2009, p. 1) Short’s
bladderpod occurrences are found in
forested sites where the canopy has
remained relatively open over time.
Common woody species associated with
Short’s bladderpod are Acer negundo
(box elder), Acer rubrum (red maple),
Aesculus glabra (Ohio buckeye), Celtis
laevigata (hackberry), Cercis canadensis
(redbud), Fraxinus Americana (white
ash), Juniperus virginiana (eastern red
cedar), Lonicera japonica (Japanese
honey suckle), Parthenocissus
quinquefolia (Virginia creeper),
Symphoricarpos orbiculatus (coral
berry) and Ulmus americana (American
elm). Common herbaceous associates
include Alliaria petiolata (garlic
mustard), Camassia scilloides (wild
hyacinth), Chaerophyllum procumbens
(spreading chervil), Delphinium tricorne
(dwarf larkspur), Galium aparine
(cleavers), Lamium sp. (dead nettle),
Phacelia bipinnatifida (forest phacelia),
Polygonatum biflorum (Solomon’s seal),
Sedum pulchellum (stonecrop), Silene
virginica (fire-pink), and Verbascum
thapsus (common mullein) (Shea 1993,
p. 19).
Biology. Published literature on the
biology of Short’s bladderpod is lacking.
The species flowers during April and
May (Gleason and Chronquist 1991, p.
187, Shea 1993, p. 20). Dr. Carol Baskin
(Professor, University of Kentucky, pers.
comm., December 2012) observed low
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fruit set in the Indiana population and,
based on lack of seed production from
plants in a greenhouse from which
pollinators were excluded, she
concluded that the species likely is selfincompatible. Self-incompatibility has
been reported in other species of
Physaria (Tepedino et al. 2012, p. 142;
Edens-Meier et al. 2011, p. 292;
Claerbout et al. 2007, p. 134; Bateman
1955, p. 64), and the molecular
mechanisms underlying self-recognition
between pollen and stigma and
subsequent pollen rejection have been
well studied in the Brassicaceae
(Takayama and Isogai 2005, pp. 468–
474). Dr. Baskin (pers. comm., December
2012) also observed that seeds produced
by Short’s bladderpod apparently are
capable of forming a seed bank, as seeds
that were planted in a greenhouse were
observed to germinate and produce
seedlings over several years, rather than
all germinating in the year they were
planted.
The pollinators for Short’s bladderpod
have not been studied, but Rollins and
Shaw (1973, p. 6) reported that bees and
flies were repeatedly observed visiting
flowers of other congeners. The majority
of floral foragers observed visiting
Physaria filiformis (Missouri
bladderpod) were true bees representing
five families, with greater than 50
percent from the family Halictidae. The
families Apidae and Andrenidae also
were well represented among bee
pollinators of this species, the most
dependable and frequent of which were
ground-nesters. Several flies of the
family Syrphidae also carried Missouri
bladderpod pollen (Edens-Meier et al.
2011, pp. 293). Tepedino et al. (2012,
pp. 143–145) found that native groundnesting bees from the families
Andrenidae and Halictidae were the
most reliable pollinators visiting flowers
of three Physaria species, but they
reported fewer numbers of pollencarrying flies from the families
Tachinidae and Conopidae. They
estimated that maximum flight distance
ranged from 100 to 1400 meters (m) (330
to 4593 feet (ft)) for the Andrenids and
40 to 100 m (130 to 330 ft) for the
Halictid bees they collected.
Whorled Sunflower
Helianthus verticillatus is a member
of the sunflower family known from
Cherokee County, Alabama; Floyd
County, Georgia; and McNairy and
Madison Counties, Tennessee. It is a
perennial arising from horizontal,
tuberous-thickened roots with slender
rhizomes. The stems are slender, erect,
and up to 2 meters (m) (6 feet (ft)) tall.
The leaves are opposite on the lower
stem, verticillate (whorled) in groups of
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3 to 4 at the mid-stem, and alternate or
opposite in the inflorescence at the end.
Individual leaves are firm in texture and
have a prominent mid-vein, but lack
prominent lateral veins found in many
members of the genus. The leaves are
linear-lanceolate in shape, narrowing at
the tip to a point, and 7.5 to 18.5 cm (3.0
to 7.2 in.) long and 0.7 to 2.0 cm (0.3
to 0.8 in.) wide. The flowers are
arranged in a branched inflorescence
typically consisting of 3 to 7 heads. The
heads are about 1 cm high (0.4 in.), are
about 1.5 cm (0.6 in.) wide, and have
deep yellow ray flowers and lighter
yellow disk flowers. The seeds are 0.4
to 0.5 cm (0.16 to 0.2 in.) long.
Several members of the aster family
are similar in appearance to whorled
sunflower, with minor morphological
differences being apparent. Helianthus
grosseserratus is similar to whorled
sunflower but its leaves typically are
arranged in an alternating pattern,
which differs from the whorled
arrangement of H. verticillatus.
Helianthus angustifolius can be
confused with H. verticillatus but it has
narrower leaves and reddish disk
flowers, as opposed to the yellow disk
flowers of H. verticillatus (Schotz 2001,
p. 1). Helianthus giganteus often
exhibits whorled leaves, but H.
verticillatus leaves have only the
midvein prominent while H. giganteus
has lateral veins evident on the leaves
(Matthews et al. 2002, p. 22).
Taxonomy. Whorled sunflower was
described by J.K. Small (1898, p. 479),
based on a collection by S.M. Bain from
Chester County, Tennessee, in 1892.
Small distinguished it from the related
H. giganteus by its smooth and hairless
stems; narrow, entire leaf blades; and
narrowly linear-lanceolate involucre (a
collection or rosette of bracts subtending
a flower cluster, umbel, or the like)
bracts (a leaflike or scalelike plant part,
usually small, sometimes showy or
brightly colored, and located just below
a flower, a flower stalk, or an
inflorescence). No additional collections
of this species had been made when
Beatley (1963, p. 153) speculated that
the specimens (which lacked basal parts
and mature seeds) from this single
collection site perhaps represented a
single aberrant individual formed from
hybridization of an opposite- and
alternate-leaved Helianthus species.
With no new material to examine,
Heiser et al. (1969, p. 209) and
Cronquist (1980, p. 36) accepted
Beatley’s suggestion that whorled
sunflower was a hybrid.
The rediscovery of the species in
1994, in Georgia, provided ample
material for reexamination of this
species’ taxonomic status. Plants
throughout these new populations were
found to conform to the morphology of
the type collection of whorled
sunflower. Morphological studies and
root-tip chromosome counts by
Matthews et al. (2002, pp. 17–23)
validated this taxon’s status as a
distinct, diploid species. The taxonomic
validity of this species was also
confirmed through genetic studies by
Ellis et al. (2006, pp. 2345–2355). Their
47115
studies showed through comparative
genetic studies with its putative parents,
H. grosseserratus and H. angustifolius,
that whorled sunflower is a good
taxonomic species of non-hybrid origin
(Ellis et al. 2006, pp. 2351–2352).
Distribution and Status. There are
four whorled sunflower populations
known to be extant, each consisting of
multiple tracked subpopulations (Table
3) (Alabama Natural Heritage Program
(ANHP) 2012, Georgia Department of
Natural Resources (GDNR), TNHID
2012). In Floyd County, Georgia, there is
one population comprised of four
subpopulations. There is one population
in Cherokee County, Alabama,
comprised of two subpopulations.
Populations in Georgia and Alabama are
less than 2 km (1.2 mi) apart. In
Tennessee, there is one population
comprised of six subpopulations in
McNairy County and one population
comprised of four subpopulations in
Madison County. Table 3 lists these
populations and subpopulations, and
relates them to EO numbers used by
State conservation agencies to track
their status. The population in Floyd
County, Georgia, is located on lands
owned by The Campbell Group, a
timber investment management
organization. This site is referred to as
the Coosa Valley Prairie and is protected
by a conservation easement held by The
Nature Conservancy, which jointly
manages the property with The
Campbell Group. All other sites also are
on private lands but are not protected.
TABLE 3—LIST OF WHORLED SUNFLOWER POPULATIONS AND SUBPOPULATIONS BY STATE AND COUNTY, WITH CORRESPONDING SITE NAMES AND ELEMENT OCCURRENCE (EO) NUMBERS FROM STATE CONSERVATION AGENCY DATABASES IN ALABAMA, GEORGIA, AND TENNESSEE
Population
(County, State)
Subpopulation
number(s)
Cherokee, AL ..................................................................
Floyd, GA ........................................................................
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Madison, TN ...................................................................
McNairy, TN ....................................................................
Status surveys have been conducted
for this species throughout its range
(Nordman 1998, pp. 1–17; 1999, pp. 1–
5; Schotz 2001, pp. 1–14; Allison 2002,
pp. 1–2; Lincicome 2003, pp. 1–2).
Despite these extensive surveys, the
number of known populations remains
low. Schotz (2001, pp. 1, 10) located 1
new population out of 44 attempts,
representing a success rate of only 2
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Site name
1
2
1
2
3
4
1–6
1–4
Kanady Creek Prairie .....................................................
Locust Branch Prairie .....................................................
Jefferson Road Wet Prairie ............................................
Kanady Creek Wet Prairie .............................................
Upper Mud Creek Wet Prairies ......................................
Sunnybell Prairie ............................................................
Turk Creek ......................................................................
Prairie Branch .................................................................
percent. Surveys during 2000 and 2002
in Tennessee were unsuccessful at
locating any additional sites (Lincicome
2003, pp. 1–2). Surveys in 2006 resulted
in discovery of the population in
McNairy County, Tennessee (Tennessee
Division of Natural Areas 2008, p. 2).
Initial efforts to estimate population
sizes of whorled sunflower relied on
counting individual stems (Allison
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Heritage
EO
Number
AL_1
AL_2
GA_1
GA_4
GA_5
GA_7
TN_2
TN_3
2002, pp. 3–8; Schotz 2001, pp. 8–10);
however, due to the species’ clonal
growth habit, stem counts overestimate
the true number of genetically distinct
individuals (genets). Ellis et al. (2006, p.
2349) found that the genetic population
size is much smaller than the number of
stems in a population and that a more
accurate population census could be
made at most whorled sunflower sites
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by counting obvious clusters of stems
rather than individual stems. However,
Mandel (2010, p. 2056) reported that
individual clusters were much less
distinct in a portion of the Alabama site
she sampled.
Ellis et al. (2006, p. 2351) counted 70
distinct clusters at the site in Madison,
Tennessee, which closely equated to 70
separate individuals through genetic
analyses; however, not all clusters were
sampled at this site (Mandel, pers.
comm., 2012). At the McNairy County,
Tennessee, population, 36 clusters of
plants were found growing along creek
banks at the unplowed edges of
cultivated crop fields and extending
into a railroad right-of-way (Tennessee
Division of Natural Areas 2008, p. 3).
Mandel (2010, p. 2056) sampled 19
clusters at the McNairy County
population and determined these
represented 24 genets; however, only
two of the four subpopulations mapped
at this population were sampled
(Mandel, pers. comm., 2012).
Mandel (2010, p. 2058) sampled the
Alabama subpopulation number 1
(Table 3) using two methods. In one
portion of the site, leaf tissue was
collected from 15 distinct clusters,
which represented 24 genets. However,
because distinct clusters were not
obvious in another portion of this
subpopulation, Mandel (2010, p. 2058)
sampled leaves from the first 100 stalks
encountered in a 1-meter-wide transect
run through the largest patch of whorled
sunflower in that area. These 100 stalks
were within an approximately 11-m (40ft) long portion of this transect, and
represented 46 distinct genets. Mandel
(2010, p. 58) estimated that 400 stalks
were present in this area and that the
total number of genets was between 100
and 200. However, more recently only
79 stems, distributed among 8 clusters,
were found at this site (Alabama Natural
Heritage Program 2011, p. 11).
Mandel (2010, p. 2056) sampled 15
clusters growing in a ‘‘wet prairie’’ at
the Georgia site, presumably
representing EO number 1 from the
Georgia Natural Heritage Program
database (Table 3). It was determined
that these clusters represented 18 genets
(Mandel 2010, p. 2058), but apparently
the other three subpopulations present
at this population were not sampled.
The true number of genets at this site is
likely much greater, as others have
reported vigorous growth of whorled
sunflower in response to prescribed
fires that are used to manage the Coosa
Valley Prairie conservation easement
area (M. Hodges, Georgia Director of
Stewardship, The Nature Conservancy,
pers. comm. May 2012; T. Patrick,
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Botanist, Georgia Department of Natural
Resources, pers. comm. February 2012).
Based on the work of Ellis (2006) and
Mandel (2010), summarized above, at
one time Alabama supported the largest
population with an estimated 100
individuals at the Kanady Creek Prairie
site, where whorled sunflower was first
found to occur in the State. However,
Schotz (2011, p. 11) found only 79
stems, distributed among 8 clusters, at
this site in 2011. Mandel (2010)
sampled only portions of the Georgia
and Tennessee populations, thus
underestimating their sizes. Whorled
sunflower likely is now most abundant
in Georgia due to population growth in
response to habitat management by The
Nature Conservancy and The Campbell
Group at the Coosa Valley Prairie.
Schotz estimated approximately 175 to
200 stems were present at the second
Alabama site in September 2008 (Schotz
pers. comm. 2009), but there were only
42 stems found at this site in 2011
(Schotz 2011, p. 14). No estimate of
individual plants is available for this
site.
Habitat. Whorled sunflower is found
in moist, prairie-like remnants, which in
a more natural condition exist as
openings in woodlands and adjacent to
creeks. Today, the only whorled
sunflower site where these habitat
conditions are present over a relatively
large area is located in the Coosa Valley
Prairie of northwest Georgia, where the
species occurs in prairie openings and
woodlands interspersed among lands
managed for pulpwood and timber
production. At one of the Alabama
subpopulations, whorled sunflower
occurs in a narrow, open strip of
vegetation between a roadside and
adjacent forest. The second Alabama
subpopulation occurs along a small
intermittent stream and adjacent
floodplain, in a site where an immature
hardwood forest was harvested in 1998.
Whorled sunflower and associated
prairie species responded favorably to
the timber removal, but the site was
soon converted into a loblolly pine
plantation and the planted seedlings
have grown into a young, dense stand
into which little light penetrates. As of
2012, there were few whorled sunflower
plants or prairie associates present at
this site. Known populations of this
species in Tennessee are relegated
mostly to narrow bands of habitat
between cultivated fields and creeks
and adjacent to roads and railroad
rights-of-way. The largest concentration
of plants in Tennessee is found at the
Madison County population, in a 1-ha
(2.5-ac) patch of remnant, wet prairie
habitat wedged between US Highway 45
and a railroad right-of-way.
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The Alabama and Georgia populations
are located on flat to gently rolling
uplands and along stream terraces in the
headwaters of Mud Creek, a tributary to
the Coosa River. In Tennessee, the
Madison County population occurs
along Turk Creek, a tributary to the
South Fork Forked Deer River, and in
adjacent uplands. The McNairy County
population occurs along Prairie Branch,
a headwater tributary to Muddy Creek
in the Tuscumbia River drainage.
We used the Natural Resources
Conservation Service’s Web Soil Survey
to determine the soil types on which
whorled sunflower populations occur
across its range (USDA, Web Soil
Survey, available online at https://
websoilsurvey.nrcs.usda.gov/app/
HomePage.htm, accessed on January 30,
2013). The most prevalent soils where
the species occurs in Georgia are
Conasauga, Lyerly, Townley, and
Wolftever silt loams and Dowellton silty
clay loam. The silt loam soils all formed
from weathered limestone or shale, and
occupy various land forms from broad
upland ridges to low stream terraces.
These soils share the characteristics of
being moderately well-drained to welldrained, being slightly to extremely
acid, and having low to moderate
fertility and organic matter content and
clayey subsoils (USDA 1978a, pp. 24–
54). The Dowellton silty clay loam
formed in alluvium (soil material
deposited by running water) on low
stream terraces and upland depressions
is poorly drained, is moderate in
fertility and organic content, is neutral
to strongly acid, and has a clayey
subsoil (USDA 1978a, pp. 28–29).
Alabama subpopulations inhabit the
Gaylesville silty clay loam, a deep,
poorly drained, slowly permeable soil
formed from limestone on floodplains
and depressed areas in limestone
valleys (USDA 1978b, p. 20). These soils
are strongly to extremely acid, with low
natural fertility and medium organic
content (USDA 1978b, p. 20). Conasauga
silt loams, discussed above, lay upslope
of the Gaylesville soils at the Alabama
whorled sunflower sites.
In Madison County, Tennessee, the
population is primarily found on Falaya
silt loam, which are poorly drained soils
that formed in alluvium derived from
loess (loamy soil material believed to be
deposited by wind) and are strongly to
very strongly acid (USDA 1978, p. 44).
The McNairy County, Tennessee,
population occurs on Iuka and Enville
fine sandy loam soils, both of which
occupy floodplains and are occasionally
flooded during winter and early spring
(USDA 1997, pp. 73–76).
The list of associated species in these
habitats indicates a community with
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strong prairie affinities. Dominant
grasses of the tall grass prairie are
present including Schizachyrium
scoparium (little bluestem),
Sorghastrum nutans (Indian grass),
Andropogon gerardii (big bluestem), and
Panicum virgatum (switch grass). Other
common herbaceous associates include
Bidens bipinnata (Spanish needles),
Carex cherokeensis (Cherokee sedge),
Hypericum sphaerocarpum (roundseed
St. Johnswort), Helianthus angustifolius
(swamp sunflower), Helenium
autumnale (common sneezeweed),
Lobelia cardinalis (cardinal flower),
Pycnanthemum virginianum (Virginia
mountain mint), Physostegia virginiana
(obedient plant), Saccharum giganteum
(sugarcane plumegrass), Silphium
terebinthinaceum (prairie rosinweed),
Sporobolus heterolepis (prairie
dropseed), and Symphyotrichum novaeangliae (New England aster) (Tennessee
Division of Natural Areas 2008, p. 5;
Matthews et al. 2002, p. 23; Schotz
2001, p. 3). Some of these areas are also
habitat for a number of other rare
species including Marshallia mohrii
(Mohr’s Barbara’s buttons), which is
federally listed as threatened.
Biology. There is little published
information available concerning the
biology of the whorled sunflower, and
the cause for its current rarity is not
known. Ellis et al. (2006, pp. 2349–
2350) investigated genetic diversity in
the Georgia, Alabama, and Madison
County, Tennessee, populations of
whorled sunflower and found high
levels of genetic diversity at the
population and species levels despite its
apparent rarity. They speculated that
this is indicative of a species that was
more widespread in the past and
perhaps became rare relatively recently
(Ellis et al. 2006, pp. 2351–2352).
Whorled sunflower populations
exhibited moderate levels of
differentiation based on markers that are
presumed to be selectively neutral, and
since these populations are
geographically distinct and ecological
conditions vary somewhat among them
Ellis et al. (2006, p. 2353) concluded
that they likely are as differentiated, if
not more so, at adaptive loci (the
specific location of a gene or DNA
sequence on a chromosome).
Whorled sunflower is a selfincompatible, clonal perennial and
flowers from August into October
(Matthews et al. 2002, pp. 17–20; Ellis
and McCauley 2008, p. 1837). The
species is easily cultivated and seed
germination is high in the laboratory.
Upon transplanting, this species has
been shown to reproduce rapidly from
rhizomes (a horizontal underground
stem that produces roots and shoots),
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creating dense colonies. The stems can
reach over 4 m (13 ft) in height
(Matthews et al. 2002, pp. 17–20).
Ellis and McCauley (2008, p. 1837)
investigated whether there were
differences among populations of
whorled sunflower with respect to
achene viability and germination rates
and whether those differences might
have a genetic basis. They conducted
this experiment for two generations of
plants, the second generation produced
from intra-population crosses of first
generation plants. They also explored
whether isolation of populations from
one another could have fitness
consequences, by conducting interpopulation crosses and evaluating
whether they found: (1) Evidence of
genetic rescue expressed as higher
fitness of hybrid individuals as
compared to any or all of the parental
populations; and (2) evidence of
outbreeding depression. Their study
included material from the Alabama,
Georgia, and Madison County,
Tennessee, populations. However, they
were unsuccessful in cultivating plants
from the Georgia population, where the
flower heads contained few viable
achenes, which produced low
germination rates (Ellis and McCauley
2008, pp. 1837–1838).
The number of crosses that produced
no viable achenes was higher in the
intra-population Tennessee crosses than
in any other pair of crossings. Those
achenes that were produced by first
generation Tennessee intra-population
crosses exhibited lower germination
rates than Alabama achenes, and second
generation Tennessee achenes from
intra-population crosses exhibited both
lower viability and germination rates
than the Alabama achenes. However,
survival rates of germinated achenes did
not differ among these populations in
either generation (Ellis and McCauley
2008, p. 1840). Ellis and McCauley
(2008, p. 1840) suggested three possible
mechanisms that could explain these
results, none of which are mutually
exclusive: (1) Limited mate availability
in the Tennessee population due to
limited diversity of self-incompatibility
alleles; (2) more extensive inbreeding
within the Tennessee population; or (3)
differential adaptation between the two
populations.
When Tennessee plants were crossed
with pollen from Alabama plants, the
second generation mean achene
viability and germination rates were
equal to or greater than those of
Alabama intra-population crosses or
Alabama plants crossed with pollen
from Tennessee plants. Mean achene
viability of Tennessee intra-population
second generation crosses was lower
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than all other groups and germination
rates were lower than both Alabama
intra-population crosses and Alabama
plants crossed with pollen from
Tennessee plants (Ellis and McCauley
2008, pp. 1839–1840).
Based on their results, Ellis and
McCauley (2008, p. 1841) concluded
that populations of whorled sunflower
are not interchangeable with respect to
phenotypic fitness-related characters
(i.e., achene viability and germination
rates) and suggested that the potential
exists for genetic rescue of the
Tennessee population by transplanting
either seeds or seedlings produced from
crosses between Tennessee and
Alabama plants into the Tennessee
population.
Fleshy-fruit Gladecress
Leavenworthia crassa is a glabrous
(morphological feature is smooth,
glossy, having no trichomes (bristles or
hair-like structures)) winter annual
known from Lawrence and Morgan
Counties, Alabama. It usually grows
from 10 to 30 cm (4 to 12 in) tall. The
leaves are mostly basal, forming a
rosette, and entire to very deeply,
pinnately (multiple leaflets attached in
rows along a central stem) lobed or
divided, to 8 cm (3.1 in) long. Flowers
are on elongating stems, and the petals
are approximately 0.8 to 1.5 cm (0.3 to
0.6 in.) long, obovate to spatulate, and
emarginate (notched at the tip). Flower
color is either yellow with orange or
white with yellow, usually with both
color forms intermixed in a single
population. The fruit is globe-shaped or
slightly more elongate and about 1.2 cm
(0.5 in) long with a slender beak at the
tip, which is 0.25 to 0.60 cm (0.1 to 0.24
in) in length. Seeds are dark brown,
nearly round in shape and winged.
Taxonomy. Fleshy-fruit gladecress
was described by Rollins in 1963, from
material collected in 1959, from Morgan
County, Alabama. Rollins (1963, pp. 61–
68) delineated the species into two
varieties (var. crassa and var. elongata)
based on differences in fruit length.
However, herbarium and field studies
have shown var. elongata to have
variation in fruit length within the range
of fruit lengths for var. crassa (McDaniel
and Lyons 1987, p. 2–3). Thus, the
species is treated as one taxon
throughout this document. This taxon
was brought to the attention of the
scientific community in 1957, by
venerable botanist Reed C. Rollins, who
distinguished the taxon from similar
species based on reproductive
morphology.
Fleshy-fruit gladecress’s globular to
oblong fruit with a smooth exterior
distinguishes it from another gladecress
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species, Leavenworthia alabamica
(Alabama gladecress), which has a much
more elongated linear fruit with
corrugated surfaces. Alabama gladecress
also does not usually have the yellow
and orange flower forms found mixed in
populations of fleshy-fruit gladecress
(McDaniel and Lyons 1987, p. 10).
Distribution and Status. Fleshy-fruit
gladecress is endemic to a 21-km (13mi) radius area in north central Alabama
in Lawrence and Morgan Counties
(Rollins 1963, p. 63). A 1961 record
from Lauderdale County has never been
confirmed (McDaniel and Lyons 1987,
p. 6). Surveys by Lyons (in litt. 1981 to
R. Sutter), McDaniel and Lyons (1987, p.
5–6), and Hilton (1997, p. 12) were
unsuccessful at locating a number of
historical sites for fleshy-fruit
gladecress. McDaniel and Lyons (1987)
failed to locate eight sites previously
reported by Rollins (1963, p. 63), and
Lloyd (1965) and Hilton (1997, p.12)
were unsuccessful at locating seven
sites listed in McDaniel and Lyons
(1987, p. 5–6).
Currently there are six known extant
occurrences of fleshy-fruit gladecress
documented, three each in Morgan and
Lawrence Counties, Alabama (Table 4).
One of these occurs on U.S. Forest
Service (USFS) lands, where it is
formally protected. The majority of
other sites are actively grazed, a practice
that has, for the most part, maintained
favorable growing conditions for the
species. However, adjusting grazing
patterns to take place during the
species’ dormant cycle would greatly
reduce potential mortality of
reproducing plants while maintaining
ideal habitat conditions.
TABLE 4—LOCATION, SITE NAMES AND DESCRIPTIONS, AND ELEMENT OCCURRENCE (EO) RANKS FOR KNOWN EXTANT
FLESHY-FRUIT GLADECRESS OCCURRENCES
County
Population designation
EO
Rank
Historic site description
Lawrence ............
Bluebird Glades ............................
D .....
Stover Branch Glades ..................
C .....
Indian Tomb Hollow Glade ...........
A .....
Cedar Plains South .......................
C .....
Cedar Plains North .......................
B .....
Massey Glade ...............................
C .....
Described by ALNHP in 1995 as approx.
0.2-ha (0.5-ac) site with 1200 plants; by
2009 was reduced to 600 plants.
Two subpopulations, most in pasture, 3.16
ha (7.8 ac); 2,200 to 2,500 plants; maintained by livestock management, found
in 1961.
0.46-ha (1.1-ac) site with 1,200 to 1,300
plants; discovered 1977.
0.04-ha (0.1-ac) site with 75 to 100 plants;
discovered 1968.
1.7-ha (4.2-ac) site with 5,000 to 6,000
plants; discovered 1968.
2.75-ha (6.8-ac) site with 2,300 to 2,500
plants; discovered 1961.
Morgan ................
Land ownership
Private & State ROW.
Private.
Federal—USFS.
Private.
Private.
Private.
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ALNHP is the Alabama Natural Heritage Program.
ROW is right-of-way.
The Alabama Natural Heritage
Program determines EO ranks ranging
from A to D for sites and populations of
rare species, with A indicating the
status of the EO is considered to be
excellent, B good, C marginal, and D
poor. The EO rank is based on a
combination of standardized criteria
including quality, condition, viability,
and defensibility. Hilton (1997, pp. 13–
26) developed the specific criteria for
determining EO ranks for fleshy-fruit
gladecress and its habitat. Based on
these criteria, only one of the six
occurrences is A-ranked. It consists of
an estimated 1200+ plants in a relatively
undisturbed glade (Schotz 2009, p. 10).
Of the remaining occurrences, one has
approximately 5,000 to 6,000 plants, but
is B-ranked because the site where it is
located is heavily grazed. Three
occurrences are C-ranked (2 occurrences
have approximately 2400 plants in a
degraded glade community; the other
occurrence has 75 to 100 plants but is
located in high-quality habitat), and one
is D-ranked (600 plants in a residential
area with no potential for habitat
restoration) (Schotz 2009).
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Habitat. This species is a component
of glade flora and occurs in association
with limestone outcroppings. The terms
‘‘glade’’ and ‘‘cedar glades’’ are used
interchangeably to refer to shallowsoiled, open areas that are dominated by
herbaceous plants and characterized by
exposed sheets of limestone or gravel.
Eastern red cedar (Juniperus virginiana)
trees are frequent in the deeper soils
along the edges of the glades (Hilton
1997, p. 1; Baskin et al. 1986, p. 138;
Baskin and Baskin 1985, p. 1). Glades
can vary in size from as small as a few
square meters to larger than 1 square
kilometer (km2) (0.37 square miles
(mi2)) and are characterized as having
an open, sunny aspect (lacking canopy)
(Quarterman 1950, p. 1; Rollins 1963, p.
5). Historically, glades in northern
Alabama occurred as glade complexes
where sparsely vegetated patches of
exposed, or nearly exposed, limestone
occurred in a matrix of woody
vegetation to form a mosaic of habitats
grading into one another (Hilton 1997,
pp. 1, 5, 64). Herbaceous diversity was
irregular over these complexes, affected
by changes in soil gradient and
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moisture, and the presence or absence of
a woody vegetation component. Few
undisturbed examples of this
community type remain (Hilton 1997,
pp. 5, 8; McDaniel and Lyons 1987, p.
11; Baskin and Baskin 1985, p. 1;
Rollins 1963, p. 5–6).
Populations of fleshy-fruit gladecress
are now located in glade-like remnants
exhibiting various degrees of
disturbance, including pastures,
roadside rights-of-way, and cultivated or
plowed fields (Hilton 1997, p. 5). As
with most of the cedar glade endemics,
fleshy-fruit gladecress exhibits weedy
tendencies, and it is not uncommon to
find the species growing in altered
habitats. However, none of the cedar
glade endemics appear to have spread
very far from their original glade
habitats; thus the geographic range of
fleshy fruit gladecress is probably very
similar to what it was in pre-settlement
times (Baskin et al. 1986, p. 140).
All species within the small genus
Leavenworthia are adapted to the
unique physical characteristics of glade
habitats, perhaps the most important of
these being a combination of shallow
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depth and high calcium content of soils
and their tendency to have temporarily
high moisture content at or very near
the surface (Rollins 1963, pp. 4–6).
Typically, only a few inches of soil
overlie the bedrock, or, in spots, the soil
may be almost lacking and the surface
barren. The glade habitats that support
all Leavenworthia species are extremely
wet during the late winter and early
spring, and become extremely dry in
summer (Rollins 1963, p. 5).
In northern Alabama, cedar glades
primarily are distributed within the
Moulton Valley subdivision of the
Interior Low Plateau Physiographic
Province, and a few glades are scattered
up the Eastern Valley subdivision of the
Tennessee Valley (Hilton 1997, p. 1).
Most of these glades are concentrated in
the Moulton Valley, a level area
underlain by Mississippian age
limestone stretching across Morgan,
Lawrence, Franklin, and Colbert
Counties in northwestern Alabama.
Glades occur in association with
outcrops of Bangor Limestone and
typically are level with exposed sheets
of limestone or limestone gravel
interspersed with fingers of cedarhardwood vegetation. The Bangor
Limestone underlying the Moulton
Valley tapers to an end in eastern
Morgan County, where it meets the
sandstone of Brindley Mountain.
Limestone is often near the soil surface,
and can be seen in rocky cultivated
fields and as small outcroppings at the
base of low-lying forested hills (Hilton
1997).
Biology. Fleshy-fruit gladecress is an
annual, spring-flowering member of the
mustard family (Brassicaceae). As an
annual, the seeds germinate in the fall,
overwinter as rosettes, and commence a
month-long flowering period beginning
in mid-March. The first seeds mature in
late April, and during most years the
plants dry and drop all of their seeds by
the end of May. It is unlikely that all
seeds produced in spring germinate the
next fall, but the length of dormancy in
the soil is not known (McDaniel and
Lyons 1987, p. 10); thus we do not know
whether the species is capable of
forming a seed bank. Native bees in the
families’ Andrenidae and Halictidae
(sweat bees), including the species
Halictus ligatus (sweat bee), were
observed carrying pollen from
Leavenworthia crassa (fleshy-fruit
gladecress) and L. alabamica (Alabama
gladecress) in northern Alabama (Lloyd
1965).
Summary of Factors Affecting the
Species
Section 4 of the Act (16 U.S.C. 1533),
and its implementing regulations at 50
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CFR part 424, set forth the procedures
for adding species to the Federal Lists
of Endangered and Threatened Wildlife
and Plants. Under section 4(a)(1) of the
Act, we may list a species based on any
of the following five factors: (A) The
present or threatened destruction,
modification, or curtailment of its
habitat or range; (B) overutilization for
commercial, recreational, scientific, or
educational purposes; (C) disease or
predation; (D) the inadequacy of
existing regulatory mechanisms; and (E)
other natural or manmade factors
affecting its continued existence. Listing
actions may be warranted based on any
of the above threat factors, singly or in
combination. Each of these factors is
discussed below.
Short’s Bladderpod
A. The Present or Threatened
Destruction, Modification, or
Curtailment of Its Habitat or Range
Shea (1993, pp. 22–23 and 42–92) and
Tennessee Department of Environment
and Conservation (2009, p. 1–3)
discussed several threats that have
destroyed or modified Short’s
bladderpod habitat and could cause
further habitat loss or modification in
the future. These include transportation
right-of-way construction and
maintenance; impoundments and
reservoir water level manipulation;
overstory shading due to forest
succession; competition and shading
from invasive, nonnative plant species;
trash dumping; commercial and
residential construction; and livestock
grazing. Predictions of increased
frequency, duration, and intensity of
droughts across the species’ range, and
increased flooding in the Midwest
region, could portend adverse effects for
Short’s bladderpod and its habitat. We
discuss each of these threats in greater
detail below.
Transportation Right-of-Way
Construction and Maintenance
During the status survey for this
species, Shea (1993, p. 22) observed that
Short’s bladderpod habitat at three sites
(Kentucky EO 7; Tennessee EOs 7, 14)
had been destroyed or degraded by road
construction or maintenance activities.
Neither of these Tennessee occurrences
is extant today (TNHID 2012). Shea
(1993, p. 60) observed 48 plants at
Kentucky EO 7 in 1992, but noted that
the population had been much more
extensive prior to improvements of U.S.
421. Shea (1993, p. 22) also indicated
that roadside maintenance posed a
continuing threat to the species at this
location. Although approximately 100
Short’s bladderpod plants were
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observed on a steep slope above the
road cut adjacent to Kentucky EO 7 in
2004 (KNHP 2012), no plants were
found at the base of the bluff, where 21
plants had been observed in 1992 (Shea
1993, p. 60) before the road cut had
altered the habitat. Poorly timed
mowing or indiscriminate herbicide
application along the road cut at the
base of this bluff could cause mortality
of seedlings produced there from seeds
that are dispersed from the plants on the
slope above. According to data from the
KNHP (2012), a road cut was present in
2004, and no Short’s bladderpod could
be found at Kentucky EO 2, where in
1992 Shea (1993, p. 52) observed 11
Short’s bladderpod plants and observed
no apparent threats to the population.
Much of the habitat downslope of a
road, where Tennessee EO 20 once
occurred but is no longer extant, was
found to be covered with rip rap in
2008, and the remaining habitat above
and below the road was overgrown
(TDEC 2009, p. 10). Road construction
destroyed suitable habitat around
Tennessee EO 23, and Short’s
bladderpod is no longer present at the
site (TNHID 2012). Based on these data,
five Short’s bladderpod occurrences (9
percent) have been lost to habitat
destruction or modification associated
with road construction or maintenance.
Shea (1993, p. 22) identified roadside
maintenance as a threat to 12
occurrences, including two discussed
above: Indiana EO 1; Kentucky EOs 1
through 4, 7, 19, and 23; and Tennessee
EOs 2, 4, 10, and 22. In addition,
Kentucky EO 27 is located along a
mowed roadside (KNHP 2012), and
TDEC (2009, p. 2) reported that
Tennessee EOs 3 and 15 could be
affected by roadside maintenance.
Indiana EO 1 is an extant roadside
occurrence, where the species’
persistence depends on periodic
clearing of competing vegetation and
associated soil disturbance to prevent
succession of the vegetation at the site
to a forested condition that would be
unsuitable for Short’s bladderpod
(Homoya, pers. comm., December 2012).
Nonetheless, poorly timed mowing or
indiscriminate herbicide application
could negatively affect this occurrence
by disrupting reproductive cycles or
causing direct mortality of Short’s
bladderpod plants. In total, roadside
maintenance has been identified as a
threat to 15 occurrences.
Short’s bladderpod is considered
extirpated from four of the eight sites in
Kentucky where roadside maintenance
has been identified as a threat to the
species. Neither Kentucky EO 2, lost to
road construction as discussed above,
nor EO 3 is extant. No plants were
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found at Kentucky EO 3 during searches
in 2004 and 2008; however, only a few
plants had been observed here in 1994
and earlier (KNHP 2012), and the cause
for the species’ current absence is not
known. Despite the presence of 17
Short’s bladderpod plants at Kentucky
EO 19 during 2005, none were found
during visits in 2004 and 2011 (KNHP
2012). While roadside maintenance
could have contributed to loss of this
population, observations by Kentucky
Natural Heritage Program (2012)
indicate that shading or competition
from invasive species is likely a primary
cause. Short’s bladderpod was last seen
at Kentucky EO 27 in 1993, when seven
plants were found along a mowed
roadside dominated by fescue and other
weeds (KNHP 2012). This occurrence
was determined to be extirpated during
a 2011 site visit by KNHP (2012) staff.
Short’s bladderpod remains extant at
four of the eight sites in Kentucky where
roadside maintenance has been
identified as a threat to the species.
Kentucky EO 1 is considered extant, but
only three Short’s bladderpod plants—
two in 1992, and one in 2009—have
been observed at this site since the
species was first observed there in 1975.
Kentucky EO 4 was treated as two
separate populations by Shea (1993, pp.
62–65), which are now tracked as a
single occurrence (KNHP 2012). While
some plants at the base of the cliff
where Kentucky EO 4 is located are
vulnerable to roadside mowing or
herbicide application, many of the
plants are on the cliff face and
associated ledges, and no impacts from
roadside maintenance have been
documented. Short’s bladderpod
abundance at this occurrence has ranged
from a low of approximately 56
individuals in 1998, to a high of at least
400 individuals in 2004 (KNHP 2012).
As discussed above, there were
approximately 100 plants observed
above the road cut at Kentucky EO 7,
but roadside maintenance could prevent
plants from becoming established at the
base of the road cut. Kentucky EO 23
has ranged in abundance from a low of
60 plants in 2008, to a high of at least
430 plants in 2001. In 2011, there were
more than 500 seedlings present at this
site, but no flowering plants were
observed. While this occurrence is
located near a roadside, there have been
no documented impacts from roadside
maintenance.
Short’s bladderpod is considered
extirpated from two of the seven sites in
Tennessee where roadside maintenance
has been identified as a threat to the
species. At Tennessee EO 2, TDEC
(2009, p. 5) found the habitat to be too
overgrown and Short’s bladderpod
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absent during a search in 1998, and no
plants were found during a monitoring
visit in 2008. As noted above, Short’s
bladderpod was no longer present when
TDEC (2009, p. 10) observed in 2008
that the roadside habitat at Tennessee
EO 20 had been covered with rip rap
and the remaining habitat above and
below the road was overgrown.
Short’s bladderpod remains extant at
five of the seven sites in Tennessee
where roadside maintenance has been
identified as a threat to the species.
More than 500 Short’s bladderpod
plants were found at Tennessee EO 3 in
2008 (TDEC 2009, p. 6), where Shea
(1993, p. 89) found 40 plants in 1992.
This occurrence is located along a
south-facing wooded slope, north of the
Cumberland River, but very little of its
habitat would be vulnerable to
maintenance associated with the road
right-of-way to the immediate west.
Tennessee EOs 4 and 10 are located
along a roadside approximately 0.5 km
(0.3 mi) apart, and both occurrences are
estimated to number in the hundreds to
thousands of plants (TDEC 2009, p. 6–
8). While roadside maintenance could
adversely affect plants located along the
base of the roadside bluffs on which
they occur, the majorities of these
occurrences are located on ledges and
bluff tops where roadside maintenance
would be unlikely to affect them.
Tennessee EO 15 is a small occurrence
located adjacent to a bridge, on a steep
limestone bluff overlooking the Harpeth
River. While no impacts from roadside
maintenance have been observed, no
more than 20 plants have ever been
counted at this occurrence. Biologists
from TDEC (2009, p. 11) found
approximately 35 plants at Tennessee
EO 22, where Shea (1993, p. 85) found
43 reproductive plants in 1992. No
impacts from roadside maintenance
were noted during this site visit.
Four Short’s bladderpod occurrences
(7 percent) apparently have been lost to
road construction or roadside
maintenance. While 10 of the known
extant occurrences (38 percent) are
located along roadsides, where
maintenance activities such as mowing
or herbicide application could affect
them, there have been few documented
examples of such effects. In many
roadside locations, Short’s bladderpod
occurs on steep slopes or bluffs, where
roadside maintenance would be
unlikely to affect the species unless the
road was widened, requiring alteration
or removal of the slope or bluff.
Moreover, well-timed and carefully
executed right-of-way maintenance
intended to control vegetation
encroachment could be beneficial by
reducing shading and competition.
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Nonetheless, the potential exists for
road widening projects or vegetation
management efforts along road rights-ofway to destroy or modify habitat, cause
mortality of individual plants, or
diminish reproductive output at a large
proportion of sites where the species
occurs.
There are seven extant Short’s
bladderpod occurrences, and three sites
from which the species is thought to be
extirpated, located in or adjacent to the
Old Tennessee Central Railroad right-ofway (TDEC 2009, p. 3, TNHID 2012),
portions of which are not actively used
or maintained or have been sold to other
rail companies. There were hundreds to
thousands of Short’s bladderpod plants
each at three of these occurrences
(Tennessee EOs 1, 10, and 17) when
TDEC (2009, p. 4) monitored the species
in 2008. The Nashville Area
Metropolitan Planning Organization
(NAMPO) (2010, p. 98) 2035 Regional
Transportation Plan reported that the
Old Tennessee Central Railroad, which
follows the Cumberland River and
passes through Ashland City, was found
to be the most practical alignment for a
proposed commuter rail to improve
intercity commute options between the
cities of Nashville and Clarksville,
Tennessee. While no plans have been
produced for developing this proposed
commuter rail system, the 2035
Regional Transportation Plan states that
this transportation option should be
developed by 2017 (NAMPO 2010, p.
98). Habitat modification or destruction
resulting from such development could
potentially affect 27 percent of the
known extant occurrences of the
species, including some occurrences
where the species is most abundant.
Flooding and Water Level Fluctuation
Shea (1993, pp. 22–23) and TDEC
(2009, p. 2) noted that impoundments
and artificial water level manipulation
threatened several Short’s bladderpod
occurrences. This threat might be better
characterized as flooding and water
level fluctuation, regardless of cause, as
some occurrences in free-flowing river
reaches are vulnerable to this threat. For
example, the Indiana occurrence is
located near an oxbow lake that was
created in a relict channel of the
Wabash River, and it is periodically
inundated by floodwaters from the river.
In 2011, this occurrence was subjected
to a prolonged flood that killed most of
the Short’s bladderpod plants at this
location (Homoya, pers. comm.,
November 2012). There were thousands
of seedlings present at this site in 2010,
and this flood event likely eliminated
the recruitment of most, if not all, of
those seedlings into the population. At
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least 100 plants were present at this site
in 2012 (Homoya, pers. comm.,
November 2012); however, it is not
known whether these were survivors of
the flood or new plants that had
sprouted from the seed bank.
There are seven Tennessee
occurrences that TDEC (2009, p. 2)
reported could be affected by water
level manipulation. One of these,
Tennessee EO 3, is located on a wooded
slope above the upper reaches of waters
impounded by Old Hickory Lake. There
were more than 500 plants at this
location in 2008, and the position of
Short’s bladderpod within the forested
area above the zone of routine water
level fluctuation is unlikely to be
affected by manipulation of water levels
in the lake. Shea (1993, p. 90) did not
mention water level manipulation in her
assessment of threats to this occurrence.
Tennessee EO 20, also in the upper
reaches of Old Hickory Lake, is
presumed extirpated but was likely lost
to placement of rip rap along the
roadside where it occurs, as discussed
above (please see Transportation Rightof-Way Construction and Maintenance).
Tennessee EO 12 is located on bluffs
overlooking the Cumberland River but
not within an area managed as a
reservoir or lake. Shea (1993, pp. 22–23)
was unable to find this occurrence in
1992, and concluded that flooding at the
base of the bluff was the cause. In 2008,
TDEC (2009, p. 8) found approximately
50 plants at Tennessee EO 12, but they
considered Short’s bladderpod habitat
to be vulnerable to flooding at this site
due to water level fluctuation and the
position of the plants at a low elevation
on the bluff. Tennessee EOs 24 through
27 are found in soil at the river bank or
on bedrock ledges within about 1.5 m (5
ft) of the waters of Cordell Hull
Reservoir (TNHID 2012), but, with the
exception of EO 27, no more than 10
plants have ever been counted at any of
these sites. These three occurrences are
vulnerable to the effects of water level
fluctuation, as evidenced by observed
erosion within the fluctuation zone
(TNHID 2012). Tennessee EO 27 appears
to be at little risk of habitat alteration
due to water level fluctuation, as it is
located on bluff ledges above the zone
of routine water level fluctuation.
While the threat of flooding or water
level fluctuation is present at only five
extant occurrences (19 percent), one of
these is the only Indiana population of
the species, where the species has
numbered in excess of 1,000 plants in
the past (Homoya, pers. comm.,
November 2012). The four occurrences
in Tennessee threatened by water level
fluctuation are small and vulnerable to
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extirpation from even limited habitat
alteration or inundation.
Overstory Shading
The most vigorous (Shea 1992, p. 24)
and stable (TDEC 2009, p. 1) Short’s
bladderpod occurrences are found in
locations where the canopy has
remained relatively open over time.
Overstory shading appears to have been
a factor contributing to the
disappearance of Short’s bladderpod at
three sites in Kentucky (EO numbers 9,
19, and 20) and one in Tennessee (EO
2) where Shea (1992, p. 4) observed
heavy shading as a threat to the species
in 1992. Overstory shading has been
identified as a threat to Indiana EO 1
(INHDC 2012), Kentucky EO 22 (KNHP
2012), and Tennessee EOs 10, 21, and
24 (TNHID 2012), or 19 percent of
known extant occurrences. Based on
these data, canopy shading has been
implicated as a factor contributing to the
disappearance of Short’s bladderpod
from four sites and has been identified
as a limiting factor at nearly one-fifth of
remaining extant occurrences.
Competition With Nonnative Plant
Species
Competition with or shading from
invasive, nonnative herbaceous and
shrub species are cited in notes
concerning threats in database records
for three of Kentucky’s (EO numbers 4,
11, and 18) (KNHP 2012) and five of
Tennessee’s (EO numbers 8, 10, 22, 24,
and 26) (TNHID 2012) extant Short’s
bladderpod occurrences. Homoya (pers.
comm., December 2012) also lists
invasive species among the threats
affecting the single Indiana occurrence.
The species most often mentioned by
these agencies include Lonicera
japonica (Japanese honeysuckle), L.
maackii (bush honeysuckle), Alliaria
petiolata (garlic mustard), and Bromus
tectorum (downy brome grass);
however, several other invasive,
nonnative species occur in sites where
Short’s bladderpod exists, including
Ligustrum spp. (privet), Rosa multiflora
(multiflora rose), and Glechoma
hederacea (ground ivy). Competition
with or shading from these species
adversely affects Short’s bladderpod.
While this threat has been specifically
noted at approximately one-third of
Short’s bladderpod occurrences, it likely
is more widespread among occurrences
of the species and has not been reported
in database records.
Trash Dumping
Shea (1993, p. 22) identified three
Short’s bladderpod sites at which trash
dumping posed a threat (Kentucky EOs
1 and 19, Tennessee EO 20). The species
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is no longer found at two of these sites:
Kentucky EO 19, where canopy shading
has been implicated in the species’
absence, and Tennessee EO 20, where
most of the habitat for the species has
been covered by rip-rap. While Short’s
bladderpod is presumed to be extant at
Kentucky EO 1, there was only one
plant found at this site in 2009 (KNHP
2012). The species was first collected at
this site in 1957, and despite several site
visits between then and 2009, only two
plants were seen there in 1992 (KNHP
2012). TDEC (2009, p. 3) lists trash
dumping as a general threat to Short’s
bladderpod, but provides no specific
information to support this conclusion.
Livestock Grazing
Livestock grazing historically
presented a threat to Short’s
bladderpod, but we are not aware of any
threats currently posed by this land use.
In addition to potentially causing direct
harm to or loss of individual plants,
livestock grazing on the steeply sloped
sites where Short’s bladderpod typically
occurs could increase soil erosion,
potentially uprooting individual plants
and causing loss of the soil seed bank.
Shea (1993, p. 22) identified three
Kentucky sites (EOs 9, 20, and 21) at
which livestock (goats or cows) grazing
posed a threat to Short’s bladderpod.
None of these sites support the species
today, likely due to multiple factors that
degraded the habitat at those locations.
In Tennessee, Shea (1993, p. 22)
reported that EO numbers 15 and 21
were threatened by grazing. However,
more recent data from TDEC (TNHID
2012) indicate that Short’s bladderpod
has remained relatively stable at these
sites and grazing is not listed among
threats observed at these locations.
Commercial and Residential
Construction
While TDEC (2009, p. 3) lists
commercial and residential construction
among potential threats to Short’s
bladderpod, there is little
documentation of these impacts.
Tennessee EO 31, which is based on a
single herbarium collection from 1979,
was apparently lost due to construction
activities at its location within the city
of Clarksville (TNHID 2012). The only
other reference we have found for this
particular threat was an observation by
TDEC (TNHID 2012) that an area in the
vicinity of Tennessee EO 21 had been
subdivided for residential construction
on the bluffs overlooking Old Hickory
Lake. Construction-related threats to
Short’s bladderpod could include direct
destruction of habitat and the plants
found there or the indirect effects of
habitat alteration from sediment runoff
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and encroachment of invasive,
nonnative plant species from areas
disturbed during construction.
Climate Change
Our analyses under the Act include
consideration of ongoing and projected
changes in climate. The terms ‘‘climate’’
and ‘‘climate change’’ are defined by the
Intergovernmental Panel on Climate
Change (IPCC). ‘‘Climate’’ refers to the
mean and variability of different types
of weather conditions over time, with 30
years being a typical period for such
measurements, although shorter or
longer periods also may be used (IPCC
2007a, p. 78). The term ‘‘climate
change’’ thus refers to a change in the
mean or variability of one or more
measures of climate (e.g., temperature or
precipitation) that persists for an
extended period, typically decades or
longer, whether the change is due to
natural variability, human activity, or
both (IPCC 2007, p. 78). Various types
of changes in climate can have direct or
indirect effects on species. These effects
may be positive, neutral, or negative and
they may change over time, depending
on the species and other relevant
considerations, such as the effects of
interactions of climate with other
variables (e.g., habitat fragmentation)
(IPCC 2007, pp. 8–14, 18–19). In our
analyses, we use our expert judgment to
weigh relevant information, including
uncertainty, in our consideration of
various aspects of climate change.
The Intergovernmental Panel on
Climate Change (IPCC) concluded that
evidence of warming of the climate
system is unequivocal (IPCC 2007a, p.
30). Numerous long-term climate
changes have been observed including
changes in arctic temperatures and ice,
widespread changes in precipitation
amounts, ocean salinity, wind patterns
and aspects of extreme weather
including droughts, heavy precipitation,
heat waves and the intensity of tropical
cyclones (IPCC 2007b, p. 7). While
continued change is certain, the
magnitude and rate of change is
unknown in many cases. Species that
are dependent on specialized habitat
types, are limited in distribution, or
have become restricted to the extreme
periphery of their range will be most
susceptible to the impacts of climate
change.
Estimates of the effects of climate
change using available climate models
lack the geographic precision needed to
predict the magnitude of effects at a
scale small enough to discretely apply
to the range of Short’s bladderpod.
However, data on recent trends and
predicted changes for the Southeast and
Midwest United States (Karl et al. 2009,
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pp. 111–122) provide some insight for
evaluating the potential threat of climate
change to the species. Most of the range
of Short’s bladderpod lies within the
geographic area included by Karl et al.
(2009, pp. 111–122) in their summary of
regional climate impacts affecting the
Southeast region; however, the Indiana
occurrence of the species lies in the
Midwest region, just west of its
boundary with the Southeast region.
Since 1970, the average annual
temperature across the Southeast has
increased by about 2 °F, with the
greatest increases occurring during
winter months. The geographic extent of
areas in the Southeast region affected by
moderate to severe spring and summer
drought has increased over the past
three decades by 12 and 14 percent,
respectively (Karl et al. 2009, p. 111).
These trends are expected to increase.
Rates of warming are predicted to more
than double in comparison to what the
Southeast has experienced since 1975,
with the greatest increases projected for
summer months. Depending on the
emissions scenario used for modeling
change, average temperatures are
expected to increase by 4.5 °F to 9 °F
by the 2080s (Karl et al. 2009, p. 111).
While there is considerable variability
in rainfall predictions throughout the
region, increases in evaporation of
moisture from soils and loss of water by
plants in response to warmer
temperatures are expected to contribute
to increased frequency, intensity, and
duration of drought events (Karl et al.
2009, p. 112).
Projected increases in winter and
spring rainfall for the Midwest region,
as well as predictions of more intense
rainfall events throughout the year, are
expected to lead to more frequent
flooding. Despite these projected trends,
the likelihood of drought is expected to
increase in the Midwest due to
warming-induced increases in
evapotranspiration rates and longer
intervals between precipitation events
(Karl et al. 2009, pp. 120–121).
Depending on timing and intensity of
drought events, Short’s bladderpod
could be adversely affected by increased
mortality rates, reduced reproductive
output due to loss or reduced vigor of
mature plants, and reduced rates of seed
germination and seedling recruitment.
The species’ presumed ability to form a
seed bank should provide some
resilience to drought-induced
population declines; however, multiple
droughts in successive years could
diminish this resilience and lead to the
loss of occurrences. Conversely,
increased drought frequency and
severity could alter structure of
vegetation communities in which
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Short’s bladderpod occurs by slowing
rates of forest canopy development,
increasing tree mortality, and increasing
light availability for the species, which
could stimulate recruitment from
dormant seed banks and increase vigor
of plants located in areas that are
presently well-shaded. The predicted
increase in flood frequency in the
Midwest could place the Indiana
population of the species at risk, as
evidenced by the loss of large numbers
of seedlings during a prolonged flood at
this site in 2011. While climate has
changed in recent decades in regions
where Short’s bladderpod occurs and
the rate of change likely will continue
to increase into the future, we do not
have data to determine how the habitats
where Short’s bladderpod occurs will be
affected by these changes and how the
species will respond to these changes.
Conservation Efforts To Reduce Habitat
Destruction, Modification, or
Curtailment of Its Range
There have been limited conservation
efforts directed towards reducing threats
affecting Short’s bladderpod and its
habitat. The Indiana Department of
Natural Resources acquired the single
Indiana occurrence. IDNR controls
competing vegetation by mowing along
the roadside where Short’s bladderpod
occurs and attempts to stimulate
germination and seedling recruitment
with light soil disturbance. The species
has responded positively, at least in the
short term, to this management
(Homoya, pers. comm., December 2012).
In Kentucky, a Landowner Incentive
Program grant was used to manage
vegetation structure or control invasive
species at two occurrences in 2005. The
effort to control bush honeysuckle at
Kentucky EO 19 provided only a shortterm benefit, if any, for Short’s
bladderpod, as bush honeysuckle is
again well established at this site.
During 2011, no Short’s bladderpod
plants could be found at this site, and
the occurrence is presumed extirpated.
The removal of cedar trees at Kentucky
EO 23 appears to have positively
affected habitat conditions for Short’s
bladderpod, as there were more than
500 plants, mostly seedlings, observed
at the site in 2011. The Kentucky State
Nature Preserve Commission acquired
lands to establish the Rockcress Hills
State Nature Preserve, where Kentucky
EO 22 is located and where the federally
listed endangered Braun’s rockcress
(listed as Arabis perstellata, but now
recognized as Boechera perstellata) also
occurs. As discussed above, this
occurrence is threatened by shading due
to forest canopy development. These
conservation efforts have benefited three
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extant Short’s bladderpod occurrences,
but significant habitat threats remain
across the species’ range.
Summary of Factor A
The threats to Short’s bladderpod
from habitat destruction and
modification are occurring throughout
the entire range of the species. These
threats include transportation right-ofway construction and maintenance;
flooding and water level fluctuation;
overstory shading; and competition with
nonnative plant species. The population
level impacts from these activities are
expected to continue into the future.
Trash dumping, livestock grazing, and
commercial and residential construction
have been recognized as threats to
habitat for this species, but there is little
evidence that these are significant
threats to extant occurrences.
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B. Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
There has been limited collection of
Short’s bladderpod seed for
conservation purposes. The Missouri
Botanical Garden holds seed accessions
from the Indiana occurrence, four
Kentucky occurrences (EOs 4, 18, 19,
and 28), and two Tennessee occurrences
(EOs 4 and 17). Kentucky EO 19 is no
longer extant, for reasons discussed
above, but Short’s bladderpod is still
found at all of the other occurrences
from which these accessions were
collected. Dr. Carol Baskin (pers.
comm., December 2012) collected seeds
from Indiana for research on seed
ecology. We are not aware of
commercial trade in Short’s bladderpod
at this time. Indiscriminate collecting
for scientific or other purposes could be
a threat to the species due to the low
numbers of individuals at most
occurrences, but we have no data to
indicate that indiscriminate collecting
of Short’s bladderpod has occurred. On
the contrary, collections for ex situ
conservation holdings could be an
important component of future recovery
efforts for the species.
C. Disease or Predation
We are not aware of any commercial
or scientific data indicating that disease
or predation threatens the continued
existence of Short’s bladderpod.
D. The Inadequacy of Existing
Regulatory Mechanisms
Section 4(b)(1)(A) of the Act requires
the Service to take into account ‘‘those
efforts, if any, being made by any State
or foreign nation, or any political
subdivision of a State or foreign nation,
to protect such species. . . .’’ In relation
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to Factor D under the Act, we interpret
this language to require the Service to
consider relevant Federal, State, and
tribal laws, plans, regulations, and other
such mechanisms that may minimize
any of the threats we describe in threat
analyses under the other four factors, or
otherwise enhance conservation of the
species. We give strongest weight to
statutes and their implementing
regulations and to management
direction that stems from those laws and
regulations. An example would be State
governmental actions enforced under a
State statute or constitution, or Federal
action under statute.
Having evaluated the significance of
the threat as mitigated by any such
conservation efforts, we analyze under
Factor D the extent to which existing
regulatory mechanisms are inadequate
to address the specific threats to the
species. Regulatory mechanisms, if they
exist, may reduce or eliminate the
impacts from one or more identified
threats. In this section, we review
existing State and Federal regulatory
mechanisms to determine whether they
effectively reduce or remove threats to
Short’s bladderpod.
Short’s bladderpod is listed as
endangered in Indiana, Kentucky, and
Tennessee. In Indiana this listing does
not provide legal protection for the
species; although, listed species are
given special consideration when
planning government-funded projects.
Additionally, the Indiana site is located
on land owned by the IDNR where
collection or damage to plants is
prohibited.
The Kentucky Rare Plants Recognition
Act, Kentucky Revised Statutes (KRS),
chapter 146, section 600–619, directs
the KSNPC to identify plants native to
Kentucky that are in danger of
extirpation within Kentucky and report
every 4 years to the Governor and
General Assembly on the conditions and
needs of these endangered or threatened
plants. This list of endangered or
threatened plants in Kentucky is found
in the Kentucky Administrative
Regulations, title 400, chapter 3:040.
The statute (KRS 146:600–619)
recognizes the need to develop and
maintain information regarding
distribution, population, habitat needs,
limiting factors, other biological data,
and requirements for the survival of
plants native to Kentucky. This statute
does not include any regulatory
prohibitions of activities or direct
protections for any species included in
the list. It is expressly stated in KRS
146.615 that this list of endangered or
threatened plants shall not obstruct or
hinder any development or use of
public or private land. Furthermore, the
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intent of this statute is not to ameliorate
the threats identified for the species, but
it does provide information on the
species.
The Tennessee Rare Plant Protection
and Conservation Act of 1985 (T.C.A.
11–26–201) authorizes the Tennessee
Department of Environment and
Conservation (TDEC) to, among other
things: conduct investigations on
species of rare plants throughout the
state of Tennessee; maintain a listing of
species of plants determined to be
endangered, threatened, or of special
concern within the state; and regulate
the sale or export of endangered species
via a licensing system. This act forbids
persons from knowingly uprooting,
digging, taking, removing, damaging,
destroying, possessing, or otherwise
disturbing for any purpose, any
endangered species from private or
public lands without the written
permission of the landowner, lessee, or
other person entitled to possession and
prescribes penalties for violations. The
TDEC may use the list of threatened and
special concern species when
commenting on proposed public works
projects in Tennessee, and the
department shall encourage voluntary
efforts to prevent the plants on this list
from becoming endangered species.
This authority shall not, however, be
used to interfere with, delay, or impede
any public works project.
Thus, despite the fact that Short’s
bladderpod is listed as endangered by
the states of Indiana, Kentucky, and
Tennessee, these designations confer no
guarantee of protection to the species or
its habitat, whether on privately owned
or state-owned lands, unless such
protections are voluntarily extended to
the species.
E. Other Natural or Manmade Factors
Affecting Its Continued Existence
The ability of populations to adapt to
environmental change is dependent
upon genetic variation, a property of
populations that derives from its
members possessing different forms
(i.e., alleles) of the same gene (Primack
1998, p. 283). Small populations
occurring in isolation on the landscape
can lose genetic variation due to the
potentially strong influence of genetic
drift, i.e., the random change in allele
frequency from generation to generation
(Barrett and Kohn 1991, p. 8). Smaller
populations experience greater changes
in allele frequency due to drift than do
larger populations (Allendorf and
Luikart 2007, pp. 121–122). Loss of
genetic variation due to genetic drift
heightens susceptibility of small
populations to adverse genetic effects,
including inbreeding depression and
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loss of evolutionary flexibility (Primack
1998, p. 283). Deleterious effects of loss
of genetic variation through drift have
been termed drift load, which is
expressed as a decline in mean
population performance of offspring in
small populations (Willi et al. 2005, p.
2260).
The likelihood that Short’s
bladderpod is self-incompatible
presents another threat related to small
population sizes. Genetic
incompatibility prevents selffertilization or reduces successful
breeding among closely related
individuals, which can decrease mean
fitness in small populations because of
increased probability of an encounter of
two incompatible haplotypes (specific
combination of alleles at adjacent
locations (loci) on the chromosome that
are inherited as a unit) (Willi et al. 2005,
p. 2256), which would prevent seed
production in self-incompatible plants.
In small populations, less common Shaplotypes (self-incompatibility
haplotypes) might be easily lost due to
genetic drift, reducing the number of
compatible mates within the population
(Byers and Meagher 1992, p. 356).
In self-incompatible plants of the
Brassicaceae family, when pollen and
stigma share S-haplotypes at the S-locus
(self-incompatibility locus, i.e., the
position on a chromosome occupied by
the self-incompatibility gene complex),
pollen tube development is disrupted
on the stigma of the female reproductive
system (Takayama and Isogai 2005, p.
469). The stigma is the receptive
structure of the female reproductive
system in plants, which also includes
the pistil and ovary, on which pollen
grains germinate and begin development
of the pollen tube. Pollen tube formation
is necessary for fertilization of the ovary
and subsequent seed production to
occur.
Despite the presence of such a
mechanism functioning to reduce or
eliminate reproductive output among
individuals sharing S-haplotypes, in
small populations mating is likely to
occur among individuals that possess
different S-haplotypes but are
genetically similar at other loci due to
loss of alleles from the population
through genetic drift (Byers and
Meagher 1992, p. 358). Mating between
such closely related individuals is
referred to as inbreeding. Inbreeding
rates are higher in small populations
because most or all individuals in the
population are related, and inbred
individuals generally have reduced
fitness as compared to non-inbred
individuals from the same population, a
phenomenon referred to as inbreeding
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depression (Allendorf and Luikart 2007,
p. 306).
Evidence in plants of inbreeding
depression due to small population size
is provided by Heschel and Paige (1995,
p. 128), who found that plants from
populations of Ipomopsis aggregata
(scarlet gilia) with 100 or fewer
flowering individuals produced smaller
seeds with lower rates of germination
success compared to those from
populations with more than 100
flowering individuals. Heschel and
Paige (1995, p. 131) also found that seed
sizes increased and germination success
improved in response to transfer of
pollen into each of the small
populations, which they interpreted as
evidence that the reduced fitness
observed in small populations was
attributable, in part, to inbreeding
depression.
Willi et al. (2005, pp. 2263) found
evidence of the three processes
described above (reduced crosscompatibility presumably due to lack of
compatible mates carrying different Shaplotypes, reduced fitness due to
inbreeding, and drift load due to loss of
genetic variation) simultaneously
affecting small populations of a plant,
Ranunculus reptans (creeping
buttercup). Populations with low allelic
diversity, taken as an indication of longterm small population size, had higher
inbreeding levels. Inbreeding depression
in these populations was expressed as
poor clonal performance and reduced
seed production in offspring (F1 plants)
produced by crosses between plants
with high kinship coefficients. Drift
load also was expressed as a reduction
in mean seed production of F1 plants in
long-term small populations (Willi et al.
2005, p. 2260).
In evaluating threats to Short’s
bladderpod that could arise due to small
population size, we first evaluated the
limited data available concerning
abundance at each of the occurrences
across the species’ range. This
represents a conservative classification
of small population size, as available
data typically do not discriminate
among life history stages, so the number
of reproducing individuals is typically
less than what is shown in the
abundance data in Table 1 (see
Distribution and Status for the Short’s
bladderpod, above). Less than 100
individual plants have ever been
observed at one time at 12 (46 percent)
of the extant occurrences in Kentucky
(EOs 1, 11, and 28) and Tennessee (EOs
8, 12, 15, 22, 24, 26, 27, 29, and 30). The
greatest number of plants ever observed
at the small Kentucky occurrences
ranged from 2 at EO 1 to 52 at EO 11
(KNHP 2012). At the small Tennessee
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occurrences, maximum recorded
abundance ranged from 3 clusters of
plants at EO 26 to approximately 50
plants each at EOs 8, 12, 22, 27, and 29
(TNHID 2012). These small populations
are at risk of adverse effects from
reduced genetic variation and associated
drift load, increased risk of inbreeding
depression, and reduced reproductive
output due to low availability of
genetically compatible mates. Many of
these occurrences where population
sizes are small are isolated from other
occurrences, decreasing the likelihood
that they could be naturally
reestablished via seed dispersal, in the
event that local extinction occurred.
Cumulative Effects From Factors A
through E
Where two or more threats affect
Short’s bladderpod occurrences, the
effects of those threats could interact or
be compounded, producing a
cumulative adverse effect that rises
above the incremental effect of either
threat alone. The most obvious cases in
which cumulative adverse effects would
be significant are those in which small
populations (Factor E) are affected by
threats that result in destruction or
modification of habitat (Factor A). Two
occurrences in Kentucky and six in
Tennessee where small population size
was identified as a threat also face
threats to their habitats, as discussed
under Factor A above. The vulnerability
of these occurrences to habitat
modification or destruction is
heightened by effects of small
population size discussed above,
reduced resilience to recover from acute
demographic effects of habitat
disturbances, and low potential for
recolonization due to isolation from
other occurrences.
Whorled Sunflower
A. The Present or Threatened
Destruction, Modification, or
Curtailment of Its Habitat or Range
Whorled sunflower appears to be a
narrow habitat specialist, occurring in
natural wet meadows or prairies and
calcareous barrens. Such habitats likely
were more extensive in the eastern
United States before European
settlement, subsequent fire suppression,
and conversion of habitat to cropland or
residential areas (Allison 1995, p. 7).
Today these prairie areas are not very
extensive, and they often are degraded
or have been destroyed for a number of
reasons. Most remaining prairie
vegetation in the geographic area where
whorled sunflower occurs exists as
remnants along roadside and utility
rights-of-way, where prairie-like
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conditions are artificially maintained
(Allison 1995, p. 4). Where whorled
sunflower habitat remains, it faces
threats due to indiscriminate use of
mechanical or chemical vegetation
management for industrial forestry,
right-of-way maintenance, or
agricultural purposes that could
adversely affect it. Because the species
requires well-lit habitats for its growth
and reproduction, shading and
competition due to vegetation
succession in the absence of natural or
human-caused disturbance also threaten
whorled sunflower habitat.
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Industrial Forestry Practices
Industrial forestry practices have
altered much suitable whorled
sunflower habitat in Georgia and
Alabama, and currently threaten one
known subpopulation in Alabama.
While surveying potential habitat for
additional populations, J. Allison
(Botanist, Georgia Department of
Natural Resources, pers. comm., March
1999) observed that much of this
species’ prairie habitat in Georgia had
been converted to pine plantations.
Nearly all of the Georgia subpopulations
and one of the Alabama subpopulations
of whorled sunflower are located on
lands that currently are owned by The
Campbell Group, a timberland
investment advisory firm. The Georgia
subpopulations on The Campbell
Group’s lands are protected from habitat
destruction or degradation by their
inclusion in the conservation easement
area at the Coosa Valley Prairie, which
was donated to The Nature Conservancy
by the Temple-Inland Corporation, the
former owner of these lands.
With the exception of the
conservation easement area at the Coosa
Valley Prairie, The Campbell Group
typically subsoil plows planting sites to
improve drainage and conditions for
tree root development, and uses
mechanical or chemical methods to
control competing vegetation when
preparing sites for planting pine
seedlings (J. King, Area Manager, The
Campbell Group, LLC, pers. comm.,
August 2012) on its lands in Floyd
County, Georgia, and Cherokee County,
Alabama. These practices could cause
direct mortality of whorled sunflower
plants at one of the Alabama
subpopulations and could contribute to
habitat degradation caused by shading
and competition (please see ‘‘Shading
and Competition’’ below) by improving
conditions for growth of planted pines.
During timber harvests, either to thin
(i.e., reduce density of pine trees in
order to improve growth conditions for
remaining trees) or to clearcut the stand,
whorled sunflower plants at this
subpopulation could be subjected to
indirect adverse effects from soil
disturbance or to direct mortality due to
movement of harvesting equipment.
Right-of-Way Maintenance
Incompatible maintenance activities
in transportation rights-of-way have
adversely affected the whorled
sunflower in Alabama and Tennessee,
and could affect one subpopulation in
Georgia. At one of the Alabama
subpopulations, the whorled sunflower
occurs in a narrow strip of vegetation
between a roadside and adjacent pine
forest, where it is vulnerable to
mortality or reduced vigor and
reproductive output due to
indiscriminate use of herbicides or
mowing for right-of-way maintenance.
Poorly timed mowing of this right-ofway prevented flowering and seed
production in some plants at this site in
2008; however, the Alabama
Department of Conservation and Natural
Resources, Alabama Department of
Transportation, and Cherokee County
Highway Department cooperated in
placing signs at the site to mark the
presence of whorled sunflower and to
attempt to prevent this in the future (W.
Barger, Botanist, Alabama Department
of Conservation and Natural Resources,
pers. comm., February 2009); periodic
replacement might be needed due to
vandalism or removal of the signs
(Barger, pers. comm., March 2012).
Regular coordination with parties
responsible for roadside maintenance at
this location will be necessary to avoid
future adverse effects to the whorled
sunflower from indiscriminate mowing
or herbicide application.
Plants extending onto a roadside
within a powerline right-of-way at the
Madison County, Tennessee, population
were subjected to herbicide spraying in
association with roadside and powerline
maintenance in 2004, causing
significant mortality (A. Bishop,
Botanist, TDEC, pers. comm., February
2008; D. Lincicome, Natural Heritage
Program Manager, TDEC, pers. comm.,
September 2006). Similarly, plants
extending into the railroad right-of-way
at the McNairy County, Tennessee,
population are vulnerable to adverse
effects from indiscriminate herbicide
application for railroad right-of-way
maintenance. A small cluster of plants
in one of the Georgia’s subpopulations
is located on the bank of a road adjacent
to the Coosa Valley Prairie easement
area and is not protected. These data
indicate that effects of indiscriminate
use of herbicides or mowing for
vegetation management in
transportation rights-of-way could
adversely affect the whorled sunflower
populations in Alabama and Tennessee,
as well as a small subpopulation in
Georgia.
Agricultural Practices and Land
Conversion
The whorled sunflower has not been
rediscovered at the type locality in
Tennessee despite intensive surveys of
that area (Nordman 1998, p. 1–2).
However, this record is from an 1892
collection and locality information is
vague, so it is not possible to determine
why this population has been lost. In
Tennessee, much of this species’
suitable habitat presumably has been
converted for agricultural use, as
substantial proportions of the counties
in the State where the species have been
found have been in row crop production
since 1850 (Table 5) (Waisanen and
Bliss 2002; GIS data available at
https://landcover.usgs.gov/cropland,
accessed January 9, 2013). Because this
species was not seen following the
initial 1892 collection until it was
rediscovered in 1994, and was not seen
again in Tennessee until 1998, it is
impossible to know the historical
distribution and abundance of its
habitat. However, the data in Table 5
indicate that land conversion to
agricultural uses has a long and
sustained history in the Tennessee
counties where the whorled sunflower
has been found and likely has
contributed to loss of habitat and
whorled sunflower populations.
TABLE 5—PROPORTIONS OF COUNTY LAND BASE CONSIDERED IMPROVED FARMLAND FOR TENNESSEE COUNTIES WHERE
THE WHORLED SUNFLOWER HAS BEEN FOUND. REPORTED HERE FOR EACH COUNTY ARE THE HIGHEST AND LOWEST PROPORTIONS ON RECORD FOR EACH COUNTY AND THE YEARS IN WHICH THEY OCCURRED AND VALUES FOR
THE YEARS 1850 AND 1997, THE FIRST AND LAST YEARS INCLUDED IN WAISANEN AND BLISS (2002).
County
High (year)
Chester ............................................................................................................................
Madison ...........................................................................................................................
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37 (1940)
54 (1949)
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Low (year)
18 (1850)
23 (1870)
02AUP2
1850
1997
18
28
23
29
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TABLE 5—PROPORTIONS OF COUNTY LAND BASE CONSIDERED IMPROVED FARMLAND FOR TENNESSEE COUNTIES WHERE
THE WHORLED SUNFLOWER HAS BEEN FOUND. REPORTED HERE FOR EACH COUNTY ARE THE HIGHEST AND LOWEST PROPORTIONS ON RECORD FOR EACH COUNTY AND THE YEARS IN WHICH THEY OCCURRED AND VALUES FOR
THE YEARS 1850 AND 1997, THE FIRST AND LAST YEARS INCLUDED IN WAISANEN AND BLISS (2002).—Continued
County
High (year)
McNairy ............................................................................................................................
Agricultural practices, including field
preparation, herbicide use, and
harvesting of crops, are threats to both
of the known Tennessee populations,
due to the species’ presence in habitats
adjacent to actively farmed crop fields
in both locations. In July 2009, TDEC
biologists observed that one clump
consisting of two whorled sunflower
stems had been destroyed by row crop
cultivation in a previously fallow field
at the McNairy County, Tennessee,
population. Unpaved access roads
around the perimeter of this field had
also been widened, encroaching on
whorled sunflower plants (7 clumps,
140 stems) in an adjacent railroad rightof-way (Bishop, pers. comm., March
2010). With the exception of the
approximately 1-ha (2.5-ac) patch of old
field habitat discussed above (see
Habitat for the whorled sunflower,
above), the Madison County, Tennessee,
whorled sunflower population is
distributed in narrow strips of
vegetation along borders of row crop
fields and is vulnerable to mechanized
disturbance of these habitats or to
effects from herbicide application.
Based on this information we conclude
that habitat at both whorled sunflower
populations in Tennessee face
significant threats associated with
agricultural practices used in row crop
production.
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Shading and Competition
Absent natural or human-caused
disturbance, habitats where whorled
sunflower occurs are threatened by
succession of vegetation to a shrubdominated or forested condition. The
largest concentration of plants at the
Madison County, Tennessee, population
is located in a successional old field
approximately 1 ha (2.5 ac) in size,
where vegetation succession threatens
to degrade the largest patch of
contiguous habitat where the majority of
this population occurs. Woody species
present at this site include Acer
negundo (box elder), Liquidambar
styraciflua (sweetgum), and Salix nigra
(black willow) (Tennessee Division of
Natural Areas 2006, p. 5), all of which
can rapidly invade moist old field
habitats if left unmanaged. No
conservation agreements or management
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33 (1920)
Low (year)
14 (1850)
1850
1997
14
20
plans are in place to ensure that this site
receives periodic disturbance to
maintain open conditions needed for
the growth and sexual reproduction of
whorled sunflower.
The Alabama subpopulation on The
Campbell Group’s lands is located in a
site where the prior owner, TempleInland Corporation, harvested an
immature hardwood forest in 1998.
Initially this timber harvest was thought
to have adversely affected the whorled
sunflower population, but these plants
and associated prairie species
responded favorably within a few years
following the harvest. However, the site
was subsequently converted into a
loblolly pine plantation, and the trees
have attained sufficient size and density
to threaten whorled sunflower plants
due to increased shading and
competition (Schotz 2011, p. 4). As of
2012, there were few whorled sunflower
plants present at this site, and those
present were in a suppressed, vegetative
condition due to strong shading and
competition from planted pines and
vegetation growing in the understory.
Encroachment by invasive, nonnative
plants following the timber harvest and
establishment of the loblolly pine stand
also is a threat at this site (Schotz 2011,
p. 12). The second Alabama
subpopulation is relegated to a narrow
strip of vegetation between a roadside
and adjacent pine forest with a densely
vegetated understory. The spatial extent
of this subpopulation is limited by the
whorled sunflower’s inability to grow in
the shaded habitat of the adjacent forest.
Based on this information we
conclude that habitat degradation due to
shading and competition resulting from
vegetation succession currently is a
significant threat to two whorled
sunflower populations. Both of the
Alabama subpopulations and the largest
contiguous patch of suitable occupied
habitat for the species in Tennessee are
at risk from this threat.
Southeast has increased by about 2 °F,
with the greatest increases occurring
during winter months. The geographic
extent of areas in the Southeast region
affected by moderate to severe spring
and summer drought has increased over
the past three decades by 12 and 14
percent, respectively (Karl et al. 2009, p.
111). These trends are expected to
increase. Rates of warming are predicted
to more than double in comparison to
what the Southeast has experienced
since 1975, with the greatest increases
projected for summer months.
Depending on the emissions scenario
used for modeling change, average
temperatures are expected to increase by
4.5 °F to 9 °F by the 2080s (Karl et al.
2009, p. 111). While there is
considerable variability in rainfall
predictions throughout the region,
increases in evaporation of moisture
from soils and loss of water by plants in
response to warmer temperatures are
expected to contribute to increased
frequency, intensity, and duration of
drought events (Karl et al. 2009, p. 112).
The predicted increase in drought
frequency, intensity, and duration could
adversely affect the moist prairie
habitats inhabited by whorled
sunflower, by reducing soil moisture
and increasing sunflower mortality rates
or reducing flowering and seed
production rates. A positive effect of
increased drought could result from
increased mortality of woody vegetation
and reduced rates of vegetation
succession, which diminishes habitat
abundance and quality for whorled
sunflower. While climate has changed
in recent decades in the region where
whorled sunflower occurs and the rate
of change likely will continue to
increase into the future, we do not have
data to determine how the habitats
where the whorled sunflower occurs
will be affected by these changes and
how the species will respond to these
changes.
Climate Change
Conservation Efforts To Reduce Habitat
Destruction, Modification, or
Curtailment of Its Range
We discuss the topic of climate
change in greater detail above in the
Factor A threats analysis for Short’s
bladderpod, which is also applicable to
whorled sunflower. Since 1970, the
average annual temperature across the
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Temple-Inland Corporation donated a
conservation easement for the Coosa
Valley Prairie property in Georgia to
The Nature Conservancy, thereby
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02AUP2
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protecting most of the Georgia
population of this species. This site
drains into the headwaters of Mud
Creek. In 2002, The Georgia Department
of Natural Resources and The Nature
Conservancy worked with staff of
Temple-Inland to develop a 10-year
management plan for conservation of
rare species within this easement area.
Site-specific management plans for
several open wet prairies, known to
provide habitat for this species within
the easement, were developed. TempleInland implemented a prescribed burn
and selective timber harvest on 243 ha
(600 ac) of the easement in 2001, to
improve habitat conditions for whorled
sunflower and other species. TempleInland conducted additional burns
within the easement area between 2002
and 2006. Mechanical thinning and
control of invasive, exotic plants was
also a component of their management
of this site.
This easement area, now owned by
The Campbell Group, is cooperatively
managed with The Nature Conservancy
based on a jointly developed
conservation management plan, which
was revised in 2012, for the period
extending through 2016. The
management goals for the site are based
on the conservation easement and
include long-term perpetuation and
restoration of the mosaic of prairies,
woodlands, wetlands, creeks, and forest
while allowing for sustainable timber
harvesting. Protecting and enhancing
native plant communities, especially
those supporting rare species, is the
primary management objective, and
periodic timber harvesting is a
secondary objective. Portions of the tract
either have been or will be planted into
Pinus palustris (longleaf pine) as part of
the Longleaf Alliance partnership.
Prescribed fire is the primary
management tool used to perpetuate and
restore the native plant communities
and also serves silvicultural objectives.
Despite the existence of a
conservation plan and the cooperative
partnership between The Nature
Conservancy and The Campbell Group
to implement the plan, management
with prescribed fire is not a binding
condition of the conservation easement.
Thus, the potential remains that this
management could be discontinued in
the event that the property was sold to
a less cooperative landowner.
Summary of Factor A
The threats to whorled sunflower
from habitat destruction and
modification are occurring throughout
the entire range of the species. These
threats include mechanical or chemical
vegetation management associated with
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industrial forestry practices,
maintenance of transportation and
utility rights-of-way, agricultural
practices, and shading and competition.
While a conservation easement and
suitable habitat management alleviate
threats from industrial forestry that
otherwise would adversely affect the
Georgia population, one of the Alabama
whorled sunflower subpopulations
currently is threatened by industrial
forestry practices. The population-level
impacts from these activities are
expected to continue into the future.
B. Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
The whorled sunflower currently is of
limited availability in the horticultural
trade, although no negative impacts are
known to have occurred due to
collection of wild material for
commercial sale. Nonetheless, the
conspicuous, attractive flowers of this
species combined with easy access of
some sites leaves the species vulnerable
to collection or poaching. Poaching from
the small populations of whorled
sunflower that are known to exist could
contribute to altered demographic or
genetic structure of populations,
potentially diminishing their viability;
however, we have no information to
suggest this currently is an active threat
or has adversely affected populations in
the past.
C. Disease or Predation
We are not aware of any commercial
or scientific data indicating that disease
or predation threatens the continued
existence of whorled sunflower.
D. The Inadequacy of Existing
Regulatory Mechanisms
Whorled sunflower is State-listed as
endangered in Georgia and Tennessee,
but has no official State status in
Alabama. The law that provides official
protection to designated species of
plants in Georgia is known as the
Wildflower Preservation Act of 1973
(O.C.G.A. 12–6–170). Under this law, no
protected plant may be collected
without written landowner permission.
No protected plant may be transported
within Georgia without a transport tag
with a permit number affixed. Permits
are also used to regulate a wide array of
conservation activities, including plant
rescues, sale of protected species, and
propagation efforts for augmentation of
natural populations and establishment
of new ones. No protected plants may be
collected from State-owned lands
without the express permission of the
Georgia Department of Natural
Resources. The Georgia Environmental
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Policy Act (GEPA; O.C.G.A. 12–16–1),
enacted in 1991, requires that impacts to
protected species be addressed for all
projects on State-owned lands, and for
all projects undertaken by a
municipality or county if funded half or
more by State funds, or by a State grant
of more than $250,000. The provisions
of GEPA do not apply to actions of
nongovernmental entities. On private
lands, the landowner has ultimate
authority over what protection efforts, if
any, occur with regard to protected
plants (Patrick et al. 1995, p. 1 of section
titled ‘‘Legal Overview’’).
The Tennessee Rare Plant Protection
and Conservation Act of 1985 (T.C.A.
11–26–201) authorizes the Tennessee
Department of Environment and
Conservation (TDEC) to, among other
things: conduct investigations on
species of rare plants throughout the
state of Tennessee; maintain a listing of
species of plants determined to be
endangered, threatened, or of special
concern within the state; and regulate
the sale or export of endangered species
via a licensing system. This act forbids
persons from knowingly uprooting,
digging, taking, removing, damaging,
destroying, possessing, or otherwise
disturbing for any purpose, any
endangered species from private or
public lands without the written
permission of the landowner, lessee, or
other person entitled to possession and
prescribes penalties for violations. The
TDEC may use the list of threatened and
special concern species when
commenting on proposed public works
projects in Tennessee, and the
department shall encourage voluntary
efforts to prevent the plants on this list
from becoming endangered species.
This authority shall not, however, be
used to interfere with, delay, or impede
any public works project.
Thus, despite the fact that whorled
sunflower is listed as endangered by the
states of Georgia and Tennessee, these
designations confer no guarantee of
protection to the species or its habitat,
whether on privately owned or stateowned lands, unless such protections
are voluntarily extended to the species
by owners or managers of lands where
the species is present.
E. Other Natural or Manmade Factors
Affecting Its Continued Existence
The whorled sunflower is vulnerable
to localized extinction because of its
extremely restricted distribution and
small population sizes at most known
locations, which reduces the resilience
of these populations to recover from
acute demographic effects of threats to
its habitat discussed above under Factor
A. Whorled sunflower is dependent
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upon existence of prairie-like openings
or remnant roadside prairie habitats for
its survival. Alteration or elimination of
disturbance processes that maintain
these openings could result in the
extinction of populations of this species.
Further, the highly fragmented
distribution of populations within
Tennessee, combined with their
disjunct location with respect to those
in Georgia and Alabama, presumably
precludes gene flow among them and
leaves little chance of natural
recolonization of these populations in
the event of localized extinctions.
Small population size could be
affecting reproductive fitness of the
whorled sunflower. The findings of Ellis
and McCauley (2008, entire) suggest that
the Madison County, Tennessee,
population is reproductively less fit
than the Alabama population. Ellis and
McCauley (2008, p. 1840) offered two
possible explanations for reduced
reproductive fitness of the Tennessee
population, including limited mate
availability due to limited diversity of
self-incompatibility alleles, or more
extensive inbreeding. Both could be
contributing to reduced seed production
and viability rates.
Ellis and McCauley (2008, pp. 1837–
1838) could not assess the fitness of the
Georgia population because seed heads
collected for the study contained very
few viable achenes, which produced
poor germination rates. However, the
lack of viable achenes in seed heads
collected for this study suggests that
poor reproductive fitness could be a
threat in this population, as well.
emcdonald on DSK67QTVN1PROD with PROPOSALS2
Cumulative Effects From Factors A
through E
Where two or more threats affect
whorled sunflower populations, the
effects of those threats could interact or
be compounded, producing a
cumulative adverse effect that rises
above the incremental effect of either
threat alone. Cumulative adverse effects
are likely significant for whorled
sunflower because all of the populations
are small and their reproductive fitness
is likely diminished (Factor E), and the
Alabama and Tennessee populations are
affected by threats that result in
destruction or modification of habitat
(Factor A). The vulnerability of these
occurrences to habitat modification or
destruction is heightened by the effects
of small population size discussed
above, reduced resilience to recover
from acute demographic effects of these
disturbances, and low potential for
recolonization due to isolation from
other occurrences.
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Fleshy-Fruit Gladecress
Factor A. The Present or Threatened
Destruction, Modification, or
Curtailment of Its Habitat or Range
This species is endemic to cedar glade
areas in north-central Alabama that have
been significantly altered from their
original condition. More than a 50
percent loss in glade habitat has
occurred since European settlement
(Hilton 1997), with resulting glade
habitats reduced to remnants
fragmented by agriculture and
development. Hilton (1997) conducted a
thorough survey of cedar glade
communities in northern Alabama using
historical records, soil maps,
topographic maps, geology, and aerial
photography; 22 high priority glades
were identified. However, field surveys
found only five of these to be in good
condition and restorable, and only two
of these were considered high-quality
sites (Hilton, pers. comm., 1999).
Agricultural Practices
At four of the fleshy-fruit gladecress
populations, plants occur in pasture
areas, on roadside rights-of-way, and/or
in planted fields surrounded by
agriculture or residential developments
(Hilton 1997, pp. 13–27). Periodic
disturbance, such as plowing in row
crop farming, arrests succession and
maintains populations in this type of
habitat; however, plowing or herbicide
application in the spring prior to seed
set and dispersal could be detrimental
to populations. Populations are
enhanced by disturbance created from
light grazing, but heavy grazing of
pastures creates unfavorable conditions
(i.e., soil compaction, nutrient
enrichment) for fleshy-fruit gladecress.
Plants have been severely trampled
where grazing is allowed during the
height of the plant’s flowering or
fruiting period. Grazing during the
reproductive period also reduces vigor
of the populations (Schotz, 2009, p. 2).
Improving pastures with fertilizer
treatments or planting of forage grasses
could eventually result in loss of
populations due to competition. Lyons
(in litt. 1981 to R. Sutter) considered
that her failure to relocate many of the
historical fleshy-fruit gladecress sites
from the 1960s was due to the change
in agricultural practices from growing
corn to using those sites for cattle
pastures. McDaniel and Lyons (1987, p.
11) considered the trend toward
converting agricultural sites from row
crop cultivation to pasture as a primary
threat to the species.
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Transportation Right-of-Way
Maintenance
Five of the six fleshy-fruit gladecress
occurrences extend onto roadsides or
are near roads, where mowing and
herbicide application prior to seed set
pose threats to the species. Three
historical sites near roads have not been
relocated and a portion of one of the
extant populations was destroyed by
road widening and grading in the 1980s
(McDaniel and Lyons 1987, p. 7–9).
Additional road widening at this site in
recent years has further reduced the size
of this population (Schotz 2009, p. 14).
The largest population of this species
has a dirt road traversing through a
portion of the site, which has made the
site vulnerable to off-road vehicles and
dumping (Hilton 1997, p. 31). Other
sites have also been negatively affected
by trash dumping and off-road vehicles,
including the site on U.S. Forest Service
land. The U.S. Forest Service has posted
the area as closed and recently gated the
area to block all-terrain vehicle access to
the site (T. Counts, U.S. Forest Service,
in litt. 2008), which appears to have
been effective at reducing damage to the
glade (A. Cochran, U.S. Forest Service,
in litt. 2005, Schotz in litt. 2007). The
U.S. Forest Service continues to monitor
the glade site for impacts from
recreational vehicles and from other
illegal vehicle activity (A. Cochran,
pers. comm., 2011).
Shading and Competition
Winter annuals, such as fleshy-fruit
gladecress, are excluded from many
habitats because they are poor
competitors (Baskin and Baskin 1985, p.
387). As with all annuals, this species’
long-term survival at a locality is
dependent upon its ability to reproduce
and reseed there every year. Thus,
populations decline and become at risk
of local extinction if conditions remain
unsuitable for reproduction for
successive years. The most vigorous
populations of the fleshy-fruit
gladecress are located in areas which
receive full, or near full, sunlight at the
canopy level and have limited
herbaceous competition (Hilton 1997, p.
5). Rollins (1963, p. 17) documented the
loss of fleshy-fruit gladecress
individuals caused by invading grasses
in an unweeded portion of an
experimental plot, while fleshy-fruit
gladecress individuals in the handweeded part of the plot thrived. Hilton
(1997, p. 12) was unable to relocate five
populations in abandoned fields and
pastures, which McDaniel and Lyons
(1987, p. 7–9) had noted as appearing
suppressed due to competition from
invading weedy species.
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Shading and competition are potential
threats at the two largest populations of
fleshy-fruit gladecress (Hilton 1997, p.
68). One site, reported to be widely
open in 1968, is now partially shaded
due to closing of the canopy (Hilton
1997, p.18). Nonnative plants, including
Ligustrum vulgare (common privet) and
Lonicera maackii (bush honeysuckle),
are a significant threat in many glades
due to the ever present disturbances that
allow for their colonization (Hilton
1997, p. 68). Nonnative plant species
pose a threat to one population of the
fleshy-fruit gladecress, where they have
established near an unimproved road
traversing the site (Hilton 1997, p.18).
Under natural conditions, cedar
glades are edaphically (related to or
caused by particular soil conditions)
maintained through processes of
drought and erosion interacting with
other processes that disrupt
encroachment of competing vegetation.
Soils that develop on glades are easily
eroded, moving downslope or into
fractures in the substrate. The shallow
soil, exposed rock, and frequently hot,
dry summers create xeric conditions
that regulate competition and shading
from encroaching vegetation (Hilton
1997, p. 5; McDaniel and Lyons 1987, p.
6; Baskin et al. 1986, p. 138; Rollins
1963, p. 5). Historically, periodic fires
also likely played a role in maintaining
these communities (Shotz 2009, p. 1).
Extant occurrences of fleshy-fruit
gladecress are primarily located in areas
modified for human use. These habitat
modifications have either eliminated or
reduced the frequency of natural
disturbance processes, such as fire, that
would otherwise regulate encroachment
of competing vegetation.
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Residential and Industrial Development
Hilton (pers. comm., 1999) considered
residential and industrial development
that had taken place in the decade prior
to her study to be the primary threat to
cedar glade communities and the
primary reason for the loss of cedar
glade habitat. One of the six fleshy-fruit
gladecress populations is located in the
front yard of a private residence.
However, at this time, we know of no
projects that would lead to the
destruction of habitat where this species
is currently located.
Climate Change
We discuss the topic of climate
change in greater detail above in the
Factor A threats analysis for Short’s
bladderpod, which is also applicable to
the fleshy-fruit gladecress. Since, 1970,
the average annual temperature across
the Southeast has increased by about 2
°F, with the greatest increases occurring
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during the winter months. The
geographic extent of areas in the
Southeast region affected by moderate to
severe spring and summer drought has
increased over the past three decades by
12 and 14 percent, respectively (Karl et
al. 2009, p. 111). These trends are
expected to increase. Rates of warming
are predicted to more than double in
comparison to what the Southeast has
experienced since 1975, with the
greatest increases projected for summer
months. Depending on the emissions
scenario used for modeling change,
average temperatures are expected to
increase by 4.5 °F to 9 °F by the 2080s
(Karl et al. 2009. p. 111). While there is
considerable variability in rainfall
predictions throughout the region,
increases in evaporation of moisture
from soils and loss of water by plants in
response to warmer temperatures are
expected to contribute to increased
frequency, intensity, and duration of
drought events (Karl et al. 2009, p. 112).
A warmer climate with more frequent
droughts, but also extreme precipitation
events, may adversely affect fleshy-fruit
gladecress by altering the glade habitat
the species requires. Ephemeral seeps
and streams on glades provide
microhabitats important to the
distribution of the species (Hilton 1997,
p. 5). Climate change may also improve
habitat conditions for invasive plant
species and other plants (USFWS 2010,
p. 5). A positive effect of increased
drought could result from increased
mortality of woody vegetation and
reduced rates of vegetation succession.
While climate has changed in recent
decades in the region where fleshy-fruit
gladecress occurs and the rate of change
likely will continue to increase for the
foreseeable future, we are unable to
determine how the habitats where
fleshy-fruit gladecress occurs will be
affected by these changes and how the
species will respond to these changes.
Conservation Efforts to Reduce Habitat
Destruction, Modification, or
Curtailment of Its Range
The occurrence and its habitat on
William B. Bankhead National Forest
(WBNF) is protected due to its location
in a Native American cultural site and
the fact that cedar glade communities
are considered ‘‘rare communities’’ on
the WBNF and protected from
detrimental effects from agency actions
(A. Cochran, U.S. Forest Service, in litt.
2005). A thorough survey of limestone
and sandstone glades on the WBNF was
completed by Schotz in 2006. Nine
glades presently are known to occur on
WBNF, with sandstone glades
constituting the largest percentage of
glade surface area. The fleshy-fruit
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47129
gladecress inhabits Indian Tomb Hollow
Glade, the one limestone glade present
on WBNF, with a surface area of
approximately 2.7 ha (1.1 ac). WBNF
conducted treatment of the nonnative
invasive species Ligustrum sinense
(Chinese privet) on the Indian Tomb
Hollow Glade in the fall of 2009 and
summer of 2011. The U.S. Forest
Service has posted the area of the
gladecress population as closed to
access and monitors impacts to the
glade from off-road vehicles. Seeds from
the Indian Tomb Hollow Glade were
collected in May 2010, and sent to the
USDA National Center for Genetic
Resources Preservation for long-term
storage.
The Service funded a survey of cedar
glade habitats in the Moulton Valley
physiographic region of northwestern
Alabama, the major area for this habitat
type, in the late 1990s. A survey and
status update for all fleshy-fruit
gladecress populations was part of that
project. The Service recently funded
surveys to update information on all
populations of this species. All sites
were visited in 2006 and 2007, and
surveys continued into 2009 (Schotz
2009). This information will be used to
develop conservation measures needed
to protect and enhance populations.
Summary of Factor A
The threats to fleshy-fruit gladecress
from habitat destruction and
modification are occurring throughout
the entire range of the species. These
threats include agricultural conversion
or incompatible practices, maintenance
of transportation rights-of-way,
residential and industrial development,
and shading and competition.
Conservation efforts of the U.S. Forest
Service have removed threats associated
with off-road vehicle use and
encroachment of invasive species at one
site; however, maintenance of
transportation right-of-ways and use of
off-road vehicles could adversely affect
the remaining five extant populations.
The population-level impacts from these
activities are expected to continue into
the future.
Factor B. Overutilization for
Commercial, Recreational, Scientific, or
Educational Purposes
There is no information to suggest
that fleshy-fruit gladecress is collected
for commercial, recreational, or
educational purposes, and we have no
reason to believe that this factor will
become a threat to the species in the
future.
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Factor C. Disease or Predation
One occurrence was lost due to
infection by mustard rust in the early
1980s (Lyons and Antonovics 1991, p.
274; McDaniel and Lyons 1987, p. 11).
We have no data to indicate whether
this disease poses a significant longterm threat to the species generally.
There is no information regarding
predation of the species by wildlife.
Grazing is ongoing across the range of
the gladecress and occurs on portions of
all extant population sites; however,
there is no information to document that
cattle eat gladecress. No studies have
been conducted to investigate the effect
of grazing or herbivory specifically on
fleshy-fruit gladecress.
Factor D. The Inadequacy of Existing
Regulatory Mechanisms
The greatest threats to the gladecress
include loss of habitat and the plants
themselves due to actions that remove
the substrate under the populations or
that cover them up. These types of
actions have been associated with
conversion of native glades or pastures
with glades and outcrops to other land
uses and potentially herbicide
applications for the purpose of
controlling invasive plants. State and
Federal regulations that might help
conserve rare species on State highway
rights-of-way, including avoidance or
minimization of habitat destruction, as
well as regulations that would protect
plants from herbicide applications,
protect only already listed species, and
therefore do not apply to gladecress.
Likewise, no existing regulations protect
the species on privately owned land,
where most of the remnant gladecress
populations are found.
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Factor E. Other Natural or Manmade
Factors Affecting Its Continued
Existence
The fleshy-fruit gladecress is
vulnerable to localized extinction
because of the small number of
occurrences and the small population
sizes within the species’ limited range.
Small population sizes decrease the
resilience of individual fleshy-fruit
gladecress occurrences to recover from
effects of other threats affecting the
species’ habitat. There are only six
remaining flesh-fruit gladecress
occurrences, and only one of these is
protected. The loss of any occurrences
would significantly impact the species’
viability by reducing its redundancy on
the landscape, which would increase its
vulnerability to stochastic
environmental stressors and reduce the
species’ resilience to recover from
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effects of threats discussed in the above
sections.
Three of the six populations of fleshyfruit gladecress are small in size as a
result of effects of habitat loss discussed
in the above sections. The loss of
populations and reductions in
population sizes have resulted in spatial
isolation between these remnant
populations. These isolated populations
are vulnerable to extinction by
reductions in genetic variation among
the populations (Klank et al. 2012, pp.
1–2; Shotz, pers. comm., 2013). Based
on this information we conclude that
the small number of populations and
the small size of populations within the
species’ limited range are significant
threats to fleshy-fruit gladecress.
Cumulative Effects From Factors A
Through E
Where two or more threats affect
fleshy-fruit gladecress occurrences, the
effects of those threats could interact or
be compounded, producing a
cumulative adverse effect that rises
above the incremental effect of either
threat alone. Cumulative adverse effects
could be significant for fleshy-fruit
gladecress because three of the six
extant populations are small (Factor E)
and all but one of the extant occurrences
are affected by threats that result in the
destruction or modification of habitat.
The vulnerability of these occurrences
to habitat modification or destruction is
heightened by effects of small
population size discussed above,
reduced resilience to recover from acute
demographic effects of these
disturbances, and low potential for
recolonization due to isolation from
other occurrences.
Proposed Determinations
We have carefully assessed the best
scientific and commercial data available
regarding the past, present, and future
threats to Short’s bladderpod, whorled
sunflower, and fleshy-fruit gladecress.
Below we state which of the five factors
are determined to be threats to these
species and summarize the severity,
timing, and significance of those threats.
Short’s Bladderpod
The most significant threats to this
species are described under Listing
Factors A (the present or threatened
destruction, modification, or
curtailment of its habitat or range) and
E (other natural or manmade factors
affecting its continued existence). Based
on the Factor A analysis, we conclude
that the loss and degradation of habitat
represents the greatest threat to Short’s
bladderpod. Road construction has
caused the loss of habitat and all Short’s
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bladderpod plants at five occurrences in
the past, and roadside maintenance or
road widening could adversely affect
nearly 40 percent of the extant
occurrences of the species due to their
position in roadside habitats. Future
development of a commuter rail project
to improve intercity commute options
between the cities of Nashville and
Clarksville, Tennessee, could affect 27
percent of known extant occurrences,
including some locations where the
species is found in greatest abundance.
Flooding and water level fluctuations
threaten 19 percent of extant Short’s
bladderpod occurrences, most notably
the single Indiana occurrence, where the
species has been present in large
numbers but recently experienced a
reduction in numbers due to prolonged
flooding. Overstory shading due to
natural forest succession, combined
with shading and competition due to
invasive, nonnative shrubs and
herbaceous species presents the most
widespread, imminent threat to Short’s
bladderpod, and has been implicated in
the loss of several historic occurrences.
These threats are expected to continue
into the foreseeable future.
The Factor E analysis demonstrated
that Short’s bladderpod is vulnerable to
adverse effects of small population size,
including potential for reduced genetic
variation, low numbers of compatible
mates, increased likelihood of
inbreeding depression, and reduced
resilience to recover from acute
demographic effects of other threats to
the species and is habitat. Fewer than
100 plants have ever been observed at
one time at 12 (46 percent) of the 26
extant occurrences, and many of these
occurrences are isolated from other
occurrences. Existing threats may be
exacerbated by the effects of ongoing
and future climate change, especially
projected increases in temperature and
increased frequency and severity of
droughts in the Southeast and projected
increases in flooding in the Midwest.
Based on our review of the best
available scientific and commercial
information, we conclude that adverse
effects associated with small and often
isolated populations, as described in the
Factor E analysis, both alone and in
conjunction with the widespread threats
described under Factor A, constitute
significant threats to Short’s
bladderpod. As discussed under Factor
D, no regulatory mechanisms exist that
would prevent or restrict activities
described under Factor A that constitute
significant threats to the species.
Therefore, on the basis of best available
scientific and commercial information
we have determined that Short’s
bladderpod is in danger of extinction
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throughout all or a significant portion of
its range and that a proposed
determination as an endangered species
is appropriate.
Whorled Sunflower
The most significant threats to this
species are described under Listing
Factors A (the present or threatened
destruction, modification, or
curtailment of its habitat or range) and
E (other natural or manmade factors
affecting its continued existence). Based
on the Factor A analysis, we conclude
that the loss and degradation of habitat
represents the greatest threat to whorled
sunflower. Past and ongoing risk of
adverse effects from mechanical or
chemical vegetation management for
industrial forestry, right-of-way
maintenance, or agriculture is a threat to
three of the four extant populations of
this species. Modification of the
remnant prairie habitats that the species
occupies due to shading and
competition resulting from vegetation
succession also threatens these three
populations, limiting growth and
reproductive output of whorled
sunflower. These threats are expected to
continue in the foreseeable future. A
conservation easement and suitable
habitat management currently alleviates
these threats that otherwise would
adversely affect the Georgia population.
The Factor E analysis demonstrated
that whorled sunflower is vulnerable to
localized extinction because of its
extremely restricted distribution and
small population sizes at most known
locations. Small population size could
be affecting reproductive fitness of
whorled sunflower by limiting
availability of compatible mates or by
causing higher rates of inbreeding
among closely related individuals. Both
of these could be contributing to
reduced seed production and viability
rates, which limit the species’ ability to
recovery from acute demographic effects
of habitat loss or modification. The
species’ dependence on remnant prairie
habitats, which are isolated on the
landscape, limits the potential for
recolonization in the event that
localized extinction events occur.
Based on our review of the best
available scientific and commercial
information, we conclude that adverse
effects associated with extremely
restricted distribution and small and
isolated populations, as described in the
Factor E analysis, both alone and in
conjunction with the threats described
under Factor A, constitute significant
threats to whorled sunflower. As
discussed under Factor D, no regulatory
mechanisms exist that would prevent or
restrict activities described under Factor
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A that constitute significant threats to
the species. Therefore, on the basis of
best available scientific and commercial
information we have determined that
whorled sunflower is in danger of
extinction throughout all or a significant
portion of its range and that a proposed
determination as an endangered species
is appropriate.
Fleshy-fruit Gladecress
The most significant threats to this
species are described under Listing
Factors A (the present or threatened
destruction, modification, or
curtailment of its habitat or range) and
E (other natural or manmade factors
affecting its continued existence). Based
on the Factor A analysis, we conclude
that the loss and degradation of habitat
represents the greatest threat to fleshyfruit gladecress. The threats to fleshyfruit gladecress from habitat destruction
and modification are occurring
throughout the entire range of the
species. These threats include
agricultural conversion for use as
pasture or incompatible practices,
maintenance of transportation rights-ofway (including mowing and herbicide
treatment prior to seed set along
roadsides), the impacts of off-road
vehicles, dumping, residential and
industrial development, and shading
and competition. Conservation efforts of
the U.S. Forest Service have removed
threats associated with off-road vehicle
use and encroachment of invasive
species at one site; however,
maintenance of transportation right-ofways and use of off-road vehicles could
adversely affect the remaining five
extant populations.
Shading due to natural forest
succession and competition from
nonnative invasive plants presents a
significant threat to fleshy-fruit
gladecress, and has been implicated in
the loss of five historic occurrences. One
site, reported to be widely open in 1968,
is now partially shaded due to closing
of the canopy and the presence of
nonnative plants, including Ligustrum
vulgare (common privet) and Lonicera
maackii (bush honeysuckle), and these
are significant threats in many glades
due to the ever present disturbances that
allow for nonnative plant colonization.
These threats are expected to continue
into the foreseeable future.
The Factor E analysis demonstrated
that fleshy-fruit gladecress is vulnerable
to localized extinction because of the
small number of occurrences and the
small population sizes within its limited
range. Small population sizes decrease
the resilience of individual fleshy-fruit
gladecress occurrences to recover from
effects of other threats affecting its
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47131
habitat and reduce genetic variation
among populations. There are only six
remaining flesh-fruit gladecress
occurrences, and only one of these is
protected. The loss of any occurrences
would significantly impact the species’
viability by reducing its redundancy on
the landscape, which would increase its
vulnerability to stochastic
environmental stressors and reduce the
species’ resilience to recover from
effects of threats discussed in the above
sections.
Based on our review of the best
available scientific and commercial
information, we conclude that adverse
effects associated with limited
distribution and small population size,
as described in the Factor E analysis,
both alone and in conjunction with the
threats described under Factor A,
constitute significant threats to fleshyfruit gladecress. As discussed under
Factor D, no regulatory mechanisms
exist that would prevent or restrict
activities described under Factor A that
constitute significant threats to the
species. Therefore, on the basis of best
available scientific and commercial
information we have determined that
fleshy-fruit gladecress is in danger of
extinction throughout all or a significant
portion of its range and that a proposed
determination as an endangered species
is appropriate.
Significant Portion of the Range
The Act defines an endangered
species as ‘‘any species which is in
danger of extinction throughout all or a
significant portion of its range.’’ A major
part of the analysis of ‘‘significant
portion of the range’’ requires
considering whether the threats to the
species are geographically concentrated
in any way. If the threats are essentially
uniform throughout the species’ range,
then no portion is likely to warrant
further consideration. Based on the
threats to Short’s bladderpod, whorled
sunflower, and fleshy-fruit gladecress
throughout their entire known ranges,
we find that these species currently are
in danger of extinction throughout all of
their ranges, based on the severity and
scope of the threats described above. As
discussed above, these species are
proposed for listing as endangered
species, rather than threatened species,
because the threats are occurring now or
will in the near term, and their potential
impacts to the species would be severe
given the limited known distribution of
the species, the small population sizes
at many of the remaining sites, and the
small area occupied by many of these
populations, putting these species at
risk of extinction at the present time. As
these threats extend throughout their
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entire ranges, it is unnecessary to
determine if they are in danger of
extinction throughout a significant
portion of their ranges. Therefore, on the
basis of the best available scientific and
commercial data, we propose listing
Short’s bladderpod, whorled sunflower,
and fleshy-fruit gladecress as
endangered species throughout their
ranges in accordance with sections 3(6)
and 4(a)(1) of the Act.
Available Conservation Measures
Conservation measures provided to
species listed as endangered or
threatened species under the Act
include recognition, recovery actions,
requirements for Federal protection, and
prohibitions against certain practices.
Recognition through listing results in
public awareness and conservation by
Federal, State, Tribal, and local
agencies; private organizations; and
individuals. The Act encourages
cooperation with the States and requires
that recovery actions be carried out for
all listed species. The protection
required by Federal agencies and the
prohibitions against certain activities
are discussed, in part, below.
The primary purpose of the Act is the
conservation of endangered and
threatened species and the ecosystems
upon which they depend. The ultimate
goal of such conservation efforts is the
recovery of these listed species, so that
they no longer need the protective
measures of the Act. Subsection 4(f) of
the Act requires the Service to develop
and implement recovery plans for the
conservation of endangered and
threatened species. The recovery
planning process involves the
identification of actions that are
necessary to halt or reverse the species’
decline by addressing the threats to its
survival and recovery. The goal of this
process is to restore listed species to a
point where they are secure, selfsustaining, and functioning components
of their ecosystems.
Recovery planning includes the
development of a recovery outline
shortly after a species is listed and
preparation of a draft and final recovery
plan. The recovery outline guides the
immediate implementation of urgent
recovery actions and describes the
process to be used to develop a recovery
plan. Revisions of the plan may be done
to address continuing or new threats to
the species, as new substantive
information becomes available. The
recovery plan identifies site-specific
management actions that set a trigger for
review of the five factors that control
whether a species remains endangered
or may be downlisted or delisted, and
methods for monitoring recovery
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progress. Recovery plans also establish
a framework for agencies to coordinate
their recovery efforts and provide
estimates of the cost of implementing
recovery tasks. Recovery teams
(comprised of species experts, Federal
and State agencies, nongovernment
organizations, and stakeholders) are
often established to develop recovery
plans. When completed, the recovery
outline, draft recovery plan, and the
final recovery plan will be available on
our Web site (https://www.fws.gov/
endangered), or from the Service’s
Tennessee Ecological Services Field
Office (see FOR FURTHER INFORMATION
CONTACT).
Implementation of recovery actions
generally requires the participation of a
broad range of partners, including other
Federal agencies, States, Tribal,
nongovernmental organizations,
businesses, and private landowners.
Examples of recovery actions include
habitat restoration (e.g., restoration of
native vegetation), research, captive
propagation and reintroduction, and
outreach and education. The recovery of
many listed species cannot be
accomplished solely on Federal lands
because their range may occur primarily
or solely on non-Federal lands. To
achieve recovery of these species
requires cooperative conservation efforts
on private, State, and Tribal lands.
If these species are listed, funding for
recovery actions will be available from
a variety of sources, including Federal
budgets, State programs, and cost share
grants for non-Federal landowners, the
academic community, and
nongovernmental organizations. In
addition, pursuant to section 6 of the
Act, the States of Indiana, Kentucky,
and Tennessee would be eligible for
Federal funds to implement
management actions that promote the
protection or recovery of Short’s
bladderpod. The States of Georgia and
Tennessee would eligible for Federal
funds to implement management
actions that promote the protection or
recovery of whorled sunflower.
Information on our grant programs that
are available to aid species recovery can
be found at: https://www.fws.gov/grants.
Although Short’s bladderpod,
whorled sunflower, and fleshy-fruit
gladecress are only proposed for listing
under the Act at this time, please let us
know if you are interested in
participating in recovery efforts for this
species. Additionally, we invite you to
submit any new information on this
species whenever it becomes available
and any information you may have for
recovery planning purposes (see FOR
FURTHER INFORMATION CONTACT).
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Section 7(a) of the Act requires
Federal agencies to evaluate their
actions with respect to any species that
is proposed or listed as an endangered
or threatened species and with respect
to its critical habitat, if any is
designated. Regulations implementing
this interagency cooperation provision
of the Act are codified at 50 CFR part
402. Section 7(a)(4) of the Act requires
Federal agencies to confer with the
Service on any action that is likely to
jeopardize the continued existence of a
species proposed for listing or result in
destruction or adverse modification of
proposed critical habitat. If a species is
listed subsequently, section 7(a)(2) of
the Act requires Federal agencies to
ensure that activities they authorize,
fund, or carry out are not likely to
jeopardize the continued existence of
the species or destroy or adversely
modify its critical habitat. If a Federal
action may affect a listed species or its
critical habitat, the responsible Federal
agency must enter into formal
consultation with the Service.
Federal agency actions within the
species’ habitat that may require
conference or consultation or both as
described in the preceding paragraph
include federally funded or permitted
actions occurring within habitat for
Short’s bladderpod, whorled sunflower,
or fleshy-fruit gladecress (e.g.,
management and any other landscape
altering activities on Federal lands
administered by the U.S. Army Corps of
Engineers or U.S. Forest Service;
issuance of section 404 Clean Water Act
(33 U.S.C. 1251 et seq.) permits by the
U.S. Army Corps of Engineers;
construction and management of gas
pipeline and power line rights-of-way
by the Federal Energy Regulatory
Commission; construction and
maintenance of roads or highways
funded or carried out by the Federal
Highway Administration; and Federal
Emergency Management Agency-funded
actions). Also subject to consultation
would be provision of Federal funds to
State and private entities through
Federal programs such as the Service’s
Partners for Fish and Wildlife Program,
State Wildlife Grant Program, and
Federal Aid in Wildlife Restoration
Program.
The Act and its implementing
regulations set forth a series of general
prohibitions and exceptions that apply
to endangered plants. All prohibitions
of section 9(a)(2) of the Act,
implemented by 50 CFR 17.61, apply.
These prohibitions, in part, make it
illegal for any person subject to the
jurisdiction of the United States to
import or export, transport in interstate
or foreign commerce in the course of a
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commercial activity, sell or offer for sale
in interstate or foreign commerce, or
remove and reduce the species to
possession from areas under Federal
jurisdiction. In addition, for plants
listed as endangered, the Act prohibits
the malicious damage or destruction on
areas under Federal jurisdiction and the
removal, cutting, digging up, or
damaging or destroying of such plants
in knowing violation of any State law or
regulation, including State criminal
trespass law. Certain exceptions to the
prohibitions apply to agents of the
Service and State conservation agencies.
The States of Georgia, Indiana,
Kentucky, and Tennessee have
regulations authorizing the
promulgation of lists of endangered
plants; however, with the exception of
Georgia, these regulations create no
obligations on the part of landowners,
public or private, to protect State-listed
plants. The Georgia Environmental
Policy Act requires that impacts to
protected species be addressed for all
projects on State-owned lands, and for
all projects undertaken by a
municipality or county if funded half or
more by State funds, or by a State grant
of more than $250,000. The Act will,
therefore, offer additional protection to
these species.
We may issue permits to carry out
otherwise prohibited activities
involving endangered and threatened
plant species under certain
circumstances. Regulations governing
permits are codified at 50 CFR 17.62 for
endangered plants, and at 17.72 for
threatened plants. With regard to
endangered plants, a permit must be
issued for the following purposes: for
scientific purposes or to enhance the
propagation or survival of the species.
It is our policy, as published in the
Federal Register on July 1, 1994 (59 FR
34272), to identify, to the maximum
extent practicable at the time a species
is listed, those activities that would or
would not constitute a violation of
section 9 of the Act. The intent of this
policy is to increase public awareness of
the effect of a proposed listing on
proposed and ongoing activities within
the range of the species proposed for
listing. The following activities could
potentially result in a violation of
section 9 of the Act; this list is not
comprehensive:
(1) Unauthorized collecting, handling,
possessing, selling, delivering, carrying,
or transporting of Short’s bladderpod,
whorled sunflower, or fleshy-fruit
gladecress, including import or export
across State lines and international
boundaries, except for properly
documented antique specimens of these
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taxa at least 100 years old, as defined by
section 10(h)(1) of the Act;
(2) Unauthorized removal, damage, or
destruction of Short’s bladderpod or
fleshy-fruit gladecress plants from
populations located on Federal land
(lands owned by the U.S. Army Corps
of Engineers or on which they hold
easements, or U.S. Forest Service lands);
and
(3) Unauthorized removal, damage or
destruction of Short’s bladderpod,
whorled sunflower, or fleshy-fruit
gladecress plants on private land in
violation of any State regulation,
including criminal trespass.
Questions regarding whether specific
activities would constitute a violation of
section 9 of the Act should be directed
to the Service’s Tennessee Ecological
Services Field Office (see FOR FURTHER
INFORMATION CONTACT). Requests for
copies of the regulations concerning
listed species and general inquiries
regarding prohibitions and permits may
be addressed to the U.S. Fish and
Wildlife Service, 105 West Park Drive,
Suite D, Athens, GA 30606; telephone
706–613–9493; facsimile 706–613–6059.
Peer Review
In accordance with our joint policy
published in the Federal Register on
July 1, 1994 (59 FR 34270), we will seek
the expert opinions of at least three
appropriate and independent specialists
regarding this proposed rule. The
purpose of peer review is to ensure that
our listing determination for these
species is based on scientifically sound
data, assumptions, and analyses. We
have invited these peer reviewers to
comment during the public comment
period.
We will consider all comments and
information received during the
comment period on this proposed rule
during preparation of a final
rulemaking. Accordingly, the final
decision may differ from this proposal.
Public Hearings
The Act provides for one or more
public hearings on this proposal, if
requested. Requests must be received
within 45 days after the date of
publication of this proposal in the
Federal Register. Such requests must be
sent to the address shown in the FOR
FURTHER INFORMATION CONTACT section.
We will schedule public hearings on
this proposal, if any are requested, and
announce the dates, times, and places of
those hearings, as well as how to obtain
reasonable accommodations, in the
Federal Register and local newspapers
at least 15 days before the hearing.
Persons needing reasonable
accommodations to attend and
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47133
participate in a public hearing should
contact the Tennessee Ecological
Services Field Office at (931) 528–6481,
as soon as possible. To allow sufficient
time to process requests, please call no
later than one week before the hearing
date. Information regarding this
proposed rule is available in alternative
formats upon request.
Required Determinations
Clarity of the Rule
Executive Order 12866 requires each
agency to write regulations that are easy
to understand. We invite your
comments on how to make this rule
easier to understand including answers
to questions such as the following:
(1) Are the requirements in the rule
clearly stated?
(2) Does the rule contain technical
language or jargon that interferes with
its clarity?
(3) Does the format of the rule
(grouping and order of sections, use of
headings, paragraphing, etc.) aid or
reduce its clarity?
(4) Would the rule be easier to
understand if it were divided into more
(but shorter) sections?
(5) Is the description of the rule in the
SUPPLEMENTARY INFORMATION section of
the preamble helpful in understanding
the emergency rule? What else could we
do to make the rule easier to
understand?
Send a copy of any comments that
concern how we could make this rule
easier to understand to Office of
Regulatory Affairs, Department of the
Interior, Room 7229, 1849 C Street NW.,
Washington, DC 20240. You also may
email the comments to this address:
Exsec@ios.goi.gov.
National Environmental Policy Act (42
U.S.C. 4321 et seq.)
We have determined that
environmental assessments and
environmental impact statements, as
defined under the authority of the
National Environmental Policy Act of
1969, need not be prepared in
connection with listing a species as an
endangered or threatened species under
the Act. We published a notice outlining
our reasons for this determination in the
Federal Register on October 25, 1983
(48 FR 49244).
References Cited
A complete list of all references cited
in this rule is available on the Internet
at https://www.regulations.gov under
Docket No. FWS–R4–ES–2013–0087 or
upon request from the Field Supervisor,
Tennessee Ecological Services Field
Office (see FOR FURTHER INFORMATION
CONTACT section).
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Federal Register / Vol. 78, No. 149 / Friday, August 2, 2013 / Proposed Rules
Authors
The primary authors of this proposed
rule are the staff members of the
Tennessee Ecological Services Field
Office (see FOR FURTHER INFORMATION
CONTACT) and the Alabama Ecological
Services Field Office.
List of Subjects in 50 CFR Part 17
Endangered and threatened species,
Exports, Imports, Reporting and
recordkeeping requirements,
Transportation.
Proposed Regulation Promulgation
2. In § 17.12 paragraph (h), add entries
for Helianthus verticillatus,
Leavenworthia crassa, and Physaria
globosa, in alphabetical order under
FLOWERING PLANTS, to the List of
Endangered and Threatened Plants, to
read as follows:
■
Accordingly, we propose to amend
part 17, subchapter B of chapter I, title
50 of the Code of Federal Regulations,
as set forth below:
PART 17—[AMENDED]
§ 17.12
1. The authority citation for part 17
continues to read as follows:
■
*
Authority: 16 U.S.C. 1361–1407; 1531–
1544; 4201–4245, unless otherwise noted.
Endangered and threatened plants.
*
*
(h) * * *
*
Species
Historic range
Scientific name
Family
Status
When listed
Common name
*
Critical
habitat
Special
rules
FLOWERING
PLANTS
*
Helianthus
verticillatus.
*
whorled sunflower ..
*
U.S.A. (AL, GA, TN)
*
Asteraceae .............
*
E
*
....................
NA
*
Leavenworthia
crassa.
*
fleshy-fruit
gladecress.
*
U.S.A. (AL) .............
*
Brassicaceae ..........
*
E
*
....................
NA
*
Physaria globosa .....
*
Short’s bladderpod
*
U.S.A. (IN, KY, TN)
*
Brassicaceae ..........
*
E
*
....................
NA
*
*
*
*
*
*
*
*
*
*
Dated: July 18, 2013.
Stephen Guertin,
Acting Director, U.S. Fish and Wildlife
Service.
*
[FR Doc. 2013–18213 Filed 8–1–13; 8:45 am]
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*
NA
*
NA
*
NA
*
Agencies
[Federal Register Volume 78, Number 149 (Friday, August 2, 2013)]
[Proposed Rules]
[Pages 47109-47134]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2013-18213]
[[Page 47109]]
-----------------------------------------------------------------------
DEPARTMENT OF THE INTERIOR
Fish and Wildlife Service
50 CFR Part 17
[Docket No. FWS-R4-ES-2013-0087; 4500030113]
RIN 1018-AZ11
Endangered and Threatened Wildlife and Plants; Endangered Status
for Physaria globosa (Short's bladderpod), Helianthus verticillatus
(whorled sunflower), and Leavenworthia crassa (fleshy-fruit gladecress)
AGENCY: Fish and Wildlife Service, Interior.
ACTION: Proposed rule.
-----------------------------------------------------------------------
SUMMARY: We, the U.S. Fish and Wildlife Service, propose to list
Physaria globosa (Short's bladderpod), Helianthus verticillatus
(whorled sunflower), and Leavenworthia crassa (fleshy-fruit gladecress)
as endangered under the Endangered Species Act of 1973, as amended
(Act). If we finalize this rule as proposed, it would extend the Act's
protections to Physaria globosa (Short's bladderpod), Helianthus
verticillatus (whorled sunflower), and Leavenworthia crassa (fleshy-
fruit gladecress) to conserve these species.
DATES: We will accept all comments received or postmarked on or before
October 1, 2013. Comments submitted electronically using the Federal
eRulemaking Portal (see ADDRESSES section, below) must be received by
11:59 p.m. Eastern Time on the closing date. We must receive requests
for public hearings, in writing, at the address shown in the FOR
FURTHER INFORMATION CONTACT section by September 16, 2013.
ADDRESSES: You may submit comments by one of the following methods:
(1) Electronically: Go to the Federal eRulemaking Portal: https://www.regulations.gov. In the Search field, enter Docket No. FWS-R4-ES-
2013-0087, which is the docket number for this rulemaking. Then, in the
Search panel on the left side of the screen, under the Document Type
heading, click on the Proposed Rules link to locate this document. You
may submit a comment by clicking on ``Comment Now!'' If your comments
will fit in the provided comment box, please use this feature of https://www.regulations.gov, as it is most compatible with our comment review
procedures. If you attach your comments as a separate document, our
preferred file format is Microsoft Word. If you attach multiple
comments (such as form letters), our preferred format is a spreadsheet
in Microsoft Excel.
(2) By hard copy: Submit by U.S. mail or hand-delivery to: Public
Comments Processing, Attn: FWS-R4-ES-2013-0087; Division of Policy and
Directives Management; U.S. Fish and Wildlife Service; 4401 N. Fairfax
Drive, MS 2042-PDM; Arlington, VA 22203.
We request that you send comments only by the methods described
above. We will post all information received on https://www.regulations.gov. This generally means that we will post any
personal information you provide us (see the Information Requested
section below for more details).
FOR FURTHER INFORMATION CONTACT: Mary E. Jennings, Field Supervisor,
U.S. Fish and Wildlife Service, Tennessee Ecological Services Field
Office, 446 Neal Street, Cookeville, TN 38501; by telephone 931-528-
6481; or by facsimile 931-528-7075. Persons who use a
telecommunications device for the deaf (TDD) may call the Federal
Information Relay Service (FIRS) at 800-877-8339.
SUPPLEMENTARY INFORMATION:
Executive Summary
Why we need to publish a rule. Under the Act, if we intend to list
a species are endangered or threatened throughout all or a significant
portion of its range, we are required to promptly publish a proposal in
the Federal Register to list the species as endangered or threatened
and make a determination on our proposal within 1 year. Listing a
species as an endangered or threatened species can only be completed by
issuing a rule.
This rule proposes to add three plants to the Federal List of
Endangered and Threatened Plants. We are proposing to list Short's
bladderpod, whorled sunflower, and fleshy-fruit gladecress as
endangered species under the Act. Elsewhere in today's Federal
Register, we propose to designate critical habitat for the Short's
bladderpod, freshy-fruit gladecress, and the whorled sunflower.
The basis for our action. Under the Act, we may determine that a
species is an endangered or threatened species based on any of five
factors: (A) The present or threatened destruction, modification, or
curtailment of its habitat or range; (B) overutilization for
commercial, recreational, scientific, or educational purposes; (C)
disease or predation; (D) the inadequacy of existing regulatory
mechanisms; or (E) other natural or manmade factors affecting its
continued existence.
We have determined that listing is warranted for these species,
which are currently at risk throughout all of their respective ranges
due to threats related to:
For Short's bladderpod, potential future construction and
ongoing maintenance of transportation rights-of-way; prolonged
inundation and soil erosion due to flooding and water level
manipulation; overstory shading due to forest succession and shading
and competition from invasive, nonnative plant species; and small
population sizes.
For whorled sunflower, mechanical or chemical vegetation
management for industrial forestry, right-of-way maintenance, or
agriculture; shading and competition resulting from vegetation
succession; limited distribution and small population sizes.
For fleshy-fruit gladecress, loss of habitat due to
residential and industrial development; conversion of agricultural
sites for use as pasture; mowing and herbicide treatment prior to seed
production; and off-road vehicles and dumping.
We will seek peer review. We are seeking comments from
knowledgeable individuals with scientific expertise to review our
analysis of the best available science and application of that science
and to provide any additional information to improve this proposed
rule. Because we will consider all comments and information we receive
during the comment period, our final determinations may differ from
this proposal.
Information Requested
We intend that any final action resulting from this proposed rule
will be based on the best scientific and commercial data available and
be as accurate and as effective as possible. Therefore, we request
comments or information from other concerned governmental agencies,
Native American tribes, the scientific community, industry, or any
other interested parties concerning this proposed rule. We particularly
seek comments concerning:
(1) The species' biology, range, and population trends, including:
(a) Habitat requirements for feeding, reproducing, and sheltering;
(b) Genetics and taxonomy;
(c) Historical and current range, including distribution patterns;
(d) Historical and current population levels, and current and
projected trends; and
(e) Past and ongoing conservation measures for these species, their
habitats or both.
(2) The factors that are the basis for making a listing
determination for a
[[Page 47110]]
species under section 4(a) of the Act, which are:
(a) The present or threatened destruction, modification, or
curtailment of its habitat or range;
(b) Overutilization for commercial, recreational, scientific, or
educational purposes;
(c) Disease or predation;
(d) The inadequacy of existing regulatory mechanisms; or
(e) Other natural or manmade factors affecting its continued
existence.
(3) Biological, commercial trade, or other relevant data concerning
any threats (or lack thereof) to this species and regulations that may
be addressing those threats.
(4) Additional information concerning the historical and current
status, range, distribution, and population size of these species,
including the locations of any additional populations of these species.
(5) Current or planned activities in the areas occupied by these
species and possible impacts of these activities on them.
Please note that submissions merely stating support for or
opposition to the action under consideration without providing
supporting information, although noted, will not be considered in
making a determination, as section 4(b)(1)(A) of the Act directs that
determinations as to whether any species is an endangered or threatened
species must be made ``solely on the basis of the best scientific and
commercial data available.''
You may submit your comments and materials concerning this proposed
rule by one of the methods listed in the ADDRESSES section. We request
that you send comments only by the methods described in the ADDRESSES
section.
If you submit information via https://www.regulations.gov, your
entire submission--including any personal identifying information--will
be posted on the Web site. If your submission is made via a hardcopy
that includes personal identifying information, you may request at the
top of your document that we withhold this information from public
review. However, we cannot guarantee that we will be able to do so. We
will post all hardcopy submissions on https://www.regulations.gov.
Please include sufficient information with your comments to allow us to
verify any scientific or commercial information you include.
Comments and materials we receive, as well as supporting
documentation we used in preparing this proposed rule, will be
available for public inspection on https://www.regulations.gov, or by
appointment, during normal business hours, at the U.S. Fish and
Wildlife Service, Tennessee Ecological Services Field Office (see FOR
FURTHER INFORMATION CONTACT).
Background
Previous Federal Actions
The Act requires the Service to identify species of wildlife and
plants that are endangered or threatened, based on the best available
scientific and commercial data. The Act directed the Secretary of the
Smithsonian Institution to prepare a report on endangered and
threatened plant species, which was published as House Document No. 94-
51. The Service published a notice in the Federal Register on July 1,
1975 (40 FR 27824), in which we announced that more than 3,000 native
plant taxa named in the Smithsonian's report and other taxa added by
the 1975 notice would be reviewed for possible inclusion in the List of
Endangered and Threatened Plants. The 1975 notice was superseded on
December 15, 1980 (45 FR 82480), by a new comprehensive notice of
review for native plants that took into account the earlier Smithsonian
report and other accumulated information. On November 28, 1983 (48 FR
53640), a supplemental plant notice of review noted the status of
various taxa. Complete updates of the plant notice were published on
September 27, 1985 (50 FR 39526) and on February 21, 1990 (55 FR 6184).
In these reviews, Short's bladderpod (as Lesquerella globosa) was
listed as a Category 2 candidate, taxa for which information in the
possession of the Service indicated that proposing to list the species
as endangered or threatened was possibly appropriate, but for which
sufficient data on biological vulnerability and threat were not
available to support listing rules. Further biological research and
field study usually was necessary to ascertain the status of taxa in
this category.
Fleshy-fruit gladecress was recognized as consisting of two
varietal taxa in these reviews, Leavenworthia crassa var. crassa and L.
crassa var. elongata. In the 1980 review, var. crassa was listed as a
Category 2 candidate, while var. elongata was listed as a Category 1
candidate, taxa for which the Service had sufficient information to
support listing as either endangered or threatened. In the 1983, 1985,
and 1990 reviews both varieties of Leavenworthia crassa were listed as
Category 2 candidates. Many Category 2 candidate species were found not
to warrant listing, either because they were not endangered or
threatened or because they did not qualify as species under the
definitions in the Act (58 FR 51144, September 30, 1993).
In 1993, the Service eliminated candidate categories, and Short's
bladderpod and the two varieties of fleshy-fruit gladecress were no
longer candidates until they were again elevated to candidate status on
October 25, 1999 (64 FR 57534). The 1999 review elevated the species
Leavenworthia crassa (fleshy-fruit gladecress) to candidate status, but
did not recognize intraspecific taxa (varieties) due to changes in
scientifically accepted taxonomy. Whorled sunflower was first listed as
a candidate species in the 1999 review. All three of these species were
then included in subsequent candidate notices of review on October 30,
2001 (66 FR 54808), June 13, 2002 (67 FR 40657), May 4, 2004 (69 FR
24876), May 11, 2005 (70 FR 24870), September 12, 2006 (71 FR 53756),
December 6, 2007 (72 FR 69034), December 10, 2008 (73 FR 75176),
November 9, 2009 (74 FR 57804), November 10, 2010 (75 FR 69222),
October 26, 2011 (76 FR 66370), and November 21, 2012 (77 FR 69994).
Species Information
Short's bladderpod
Physaria globosa is a member of the mustard family (Brassicaceae)
known from Posey County, Indiana; Clark, Franklin and Woodford
Counties, Kentucky; and Cheatham, Davidson, Dickson, Jackson,
Montgomery, Smith, and Trousdale Counties, Tennessee. The following
description is based on Flora of North America (https://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250095135, accessed on December
7, 2012) and Gleason and Chronquist (1991, p. 187).
Short's bladderpod is an upright biennial or perennial (lives for 2
years or longer) with several stems, some branched at the base,
reaching heights up to 50 centimeters (cm) (20 inches (in.)), and which
are leafy to the base of the inflorescence (a group or cluster of
flowers arranged on a stem that is composed of a main branch or a
complicated arrangement of branches). The basal leaves, borne on short
petioles (stalks) are 2.5 to 5 cm (1 to 2 in.) in length and 0.5 to 1.5
cm (0.2 to 0.6 in.) wide, obovate (egg-shaped and flat, with the narrow
end attached to the stalk) or oblanceolate (with the widest portion of
the leaf blade beyond the middle) in shape, with a smooth or slightly
wavy margin, and gray-green in color due to a layer of dense hairs.
Leaves are gradually reduced in size and petiole length higher up the
stem. Numerous flowers are borne on a raceme (elongate,
[[Page 47111]]
spike-shaped inflorescence to which individual flowers are attached by
slender pedicels, or stalks, which in Short's bladderpod are longer
than the flowers). The yellow flowers are composed of four spoon-shaped
petals, 0.4 to 0.7 cm (0.16 to 0.28 in.) long. The fruit is globose in
shape and lightly beset with stellate (star-shaped) hairs, but becoming
smooth with time.
Taxonomy. A member of the mustard family (Brassicaceae), Short's
bladderpod was first described as Vesicaria globosa by Desvaux in 1814
(Payson 1922, pp. 103-236). Because of several distinctive characters,
Watson (1888, pp. 249-255) proposed that the American species of the
genus Vesicaria be placed in the genus Lesquerella. This treatment was
recognized as valid, until Al-Shehbaz and O'Kane (2002, entire)
reunited most of the genus Lesquerella with the genus Physaria. This
determination was supported by molecular, morphological, cytological,
biogeographic, and ecological lines of evidence (Al-Shehbaz and O'Kane
2002, p. 320). Flora of North America recognizes this change, using the
scientific name Physaria globosa for Short's bladderpod (https://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=250095135,
accessed on April 20, 2011).
Distribution and Status. In a 1992 status survey for Short's
bladderpod, Shea (1993, pp. 6-15) observed the species at only 26 of 50
historical sites: 1 in Indiana, 14 in Kentucky, and 11 in Tennessee.
The remaining sites were classified as follows (Shea 1993, p. 10-14):
Status uncertain--4 occurrences where the species had been
observed during the prior 25 years and where appropriate habitat
existed with no evidence that the occurrence had been destroyed (Shea
population numbers 27 through 30).
Extirpated--one occurrence where the habitat had been
severely altered (Shea population number 31).
Historical--5 occurrences where the species had not been
observed during the prior 25 years, but where appropriate habitat
remained (Shea population numbers 32 through 36).
Locality information incomplete--14 occurrences for which
location information was insufficient to confirm the species' presence
or absence, despite searches having been attempted in some cases (Shea
population numbers 37 through 50). Many of these putative occurrences
were based on herbarium specimens dating from the late-19th to mid-20th
centuries that contained little information about sites from which they
were collected. Except for the populations numbered 37, 42, and 50,
Shea (1993) searched for suitable habitat or Short's bladderpod plants
in areas associated with these occurrences but did not find the
species.
Later surveys found Short's bladderpod extant at two of these sites,
Tennessee element occurrence (EO) numbers 8 and 12, which correspond to
Shea's population numbers 34 and 29, respectively.
We used data provided to us by conservation agencies in the States
where the species occurs (Indiana Natural Heritage Data Center (INHDC)
2012, Kentucky Natural Heritage Program (KNHP) 2012, Tennessee
(Tennessee Natural Heritage Inventory Database (TNHID) 2012) to
determine the current distribution and status of Short's bladderpod.
Difficulty in relating the species' distribution at the time of Shea's
(1993, entire) status survey to its current distribution comes as a
result of State conservation agencies revising the mapping of some
element occurrences in these databases. In two instances, pairs of
occurrences that Shea (1993) considered distinct have been combined
into single element occurrences (Table 1). Conversely, TNHID (2012)
treats as two distinct element occurrences the two locations that Shea
(1993, p. 85, 108) mapped together as population number 23. One of
these occurrences (TN EO number 22) was extant as of 2012 (Table 1),
while the other (TN EO number 2) is extirpated (Table 2). Based on
current mapping, State conservation agencies now recognize 24 element
occurrences that correspond to populations that Shea (1993, entire)
found extant in 1992. Of these 24 occurrences, 18 were extant in 2012.
Accounting for rediscovery of the two Tennessee occurrences that Shea
(1993, pp. 10-14) did not find during 1992, and recent changes in
element occurrence mapping, a total of 20 occurrences that were
documented by Shea (1993, entire) were still considered extant as of
2012 (Table 1).
The approximate range of abundance shown in Table 1 is primarily
based on individual plants. As a result of location, it was impossible
to enumerate individual plants. This resulted in are two instances
where TNHID surveyed these populations from a boat and reported the
approximate range in clusters.
Table 1--List of Known Extant Short's Bladderpod Occurrences by State and County, With Element Occurrence (EO) Numbers Assigned by State Natural
Heritage Programs (INHDC (2012), KNHP (2012), TNHID (2012)), Numbers Assigned to Populations Reported in Shea (1993), and First and Last Years of Known
Observations
--------------------------------------------------------------------------------------------------------------------------------------------------------
EO Number
(Shea Approximate range of
State County Population First observed Last observed abundance Land ownership
Number)
--------------------------------------------------------------------------------------------------------------------------------------------------------
Indiana.......................... Posey............... 1 (1) 1941-05-06 2012 3-1000s.............. IDNR.
Kentucky......................... Clark............... 1 (3) 1957 2009-05-21 2.................... Private.
Franklin............ 4 (11, 12) 1979 2011-04-19 100-500.............. Private.
7 (10) 1981 2004-05-17 1-100................ Private.
11 (13) 1983 2003-06-01 1-52................. Private.
18 (4) 1992 2012-05-09 20-350............... City of Frankfort.
22 (9) 1990-Pres 2012-05-08 2-200................ private; Kentucky State
Nature Preserves
Commission.
23 (14) 1990 2011-04-26 60-500............... Private.
Woodford............ 28 2005-05-06 2010-06-02 few.................. Private.
Tennessee........................ Cheatham............ 1 (18) 1956-03-02 2008-04-23 100s-1000s........... COE; private.
15 (17) 1955-04-24 2008-04-29 few-20............... COE.
[[Page 47112]]
17 (16) 1953-04-26 2012-06-15 20-~1500............. Town of Ashland City;
private.
29 1998-05-12 2008-04-29 ~50.................. COE; private.
30 1998-05-12 2008-04-29 10-25................ COE; private.
Davidson; Cheatham.. 10 (21,22) 1935 2012-06-15 10s-1000s............ Private.
Davidson............ 4 (19) 1971-05-16 2012-06-15 100s-1000s........... private; COE easement.
8 (34) 1886-04-22 2008-05-02 ~50.................. private; COE easement.
Dickson............. 32 2008-04-29 2008-04-29 ~7 clusters.......... COE.
Jackson............. 26 1998-05-08 2008-05-06 3 clusters........... COE.
27 1998-05-08 2008-05-06 ~50.................. COE.
Montgomery.......... 12 (29) 1946-04-27 2008-05-09 ~50.................. private; COE easement.
22 (23a) 1969-04-28 2008-05-02 20-50................ private; COE easement.
28 1998-04-23 2008-04-29 ~300................. private; COE easement.
Smith............... 24 1998-05-05 2008-05-06 ~10.................. COE.
Trousdale........... 3 (25) 1969-05-08 2008-05-06 40-500............... COE; private.
21 (26) 1992-04-30 2008-05-12 100-250.............. COE; private.
--------------------------------------------------------------------------------------------------------------------------------------------------------
IDNR is the Indiana Department of Natural Resources.
COE is the U.S. Army Corps of Engineers.
Pres is present.
Despite the rediscovery of the two Tennessee occurrences and the
discovery of 10 additional occurrences since the 1992 status survey,
only 26 extant occurrences of Short's bladderpod are known to remain
due to the loss of 10 occurrences during the last 20 years (Table 1).
Seven of the occurrences that Shea (1993, pp. 44-71) observed in 1992,
and three others (Kentucky EO number 27 and Tennessee EO numbers 23 and
25) that were seen after 1992, have since been extirpated (Table 2).
This constitutes a loss of 27 percent of all occurrences that were
extant during 1992 or later.
Table 2--List of Extirpated Short's Bladderpod Occurrences by State and County, With Element Occurrence (EO) Numbers Assigned by State Natural Heritage
Programs (INHDC (2012), KNHP (2012), TNHID (2012)), Numbers Assigned to Populations Reported in Shea (1993), and First and Last Years of Known
Observations
--------------------------------------------------------------------------------------------------------------------------------------------------------
EO Number
(Shea
State County Population First observed Last observed Abundance Land ownership
Number)
--------------------------------------------------------------------------------------------------------------------------------------------------------
Kentucky........................... Bourbon............... * 19 (2) 1963-04-27 2005-06-09 10-120 private.
Fayette............... 12 (38) 1931 1931-05-24 n/a private.
16 (37) 1892 1900-05-09 n/a private.
Franklin.............. * 2 (6) 1979-05 1992-05-04 11 private.
* 3 (8) 1979 1994-05-12 4 private.
5 (39) 1880 1880-06 n/a private.
8 (27) 1981 1981-05-03 ~40 private.
14 (40) 1856 1856-05 n/a private.
* 20 (5) 1992 1992-05-19 21 private.
* 21 (7) 1992 1992-05-12 7 private.
Jessamine............. 6 (42) 1942 1942-05-16 n/a private.
13 (32) 1939 1939-04-27 n/a private.
17 (28) 1991-Pre 1991-Pre n/a private.
\+\ 27 1990 1993-05-10 1-7 private.
Madison............... 10 (43) 1903 1903-05-16 n/a private.
Mercer................ 24 (44) 1916 1916-05-13 1-7 private.
Nelson................ 25 1935-pre 1935-pre n/a private.
Powell................ 15 (45) 1923 1923-05-26 n/a private.
Scott................. * 9 (15) 1930 1992-05-19 2 private.
Tennessee.......................... Cheatham.............. 14 (33) 1969-04-29 1969-04-29 n/a private.
Davidson.............. * 9 (20) 1974-04-16 1998-04-16 20-29 private; COE easement.
\+\ 23 1997-05-09 1997-05-09 ~200 private.
[[Page 47113]]
Jackson............... \+\ 25 1998-07-24 1998-07-24 5 COE
Maury................. 7 (31) 1955-04-23 1955-04-23 n/a private.
Montgomery............ 2 (23b) 1968-05-07 1992-04-28 1 private.
13 (30) 1975-05-25 1975-05-25 n/a private.
18 (35) 1967-06-01 1967-06-01 n/a private.
31 1979-04-09 1979-04-09 .............. private.
Smith................. 20 (24) 1992-05-01 1998-04-17 30 private; COE easement.
--------------------------------------------------------------------------------------------------------------------------------------------------------
* Occurrences observed by Shea (1993), but which are now considered extirpated.
\+\ Occurrences not documented in Shea (1993) that have been observed since 1992, but which are now considered extirpated.
COE is the U.S. Army Corps of Engineers.
Pres is present.
No records exist in State-maintained databases for seven
populations that Shea (1993, pp. 12-13) treated as historical or
lacking sufficient locality information to verify (population number 41
from Kentucky, and numbers 36 and 46 through 50 from Tennessee).
Therefore, Table 1 and Table 2 do not include entries for these Shea
population numbers. Shea (1993, p. 15) also determined that four
historical reports for the species were erroneous: One each from Monroe
County, Indiana, and Vinton County, Ohio; and one each from unknown
counties in Kansas and Vermont.
There are now 8 known extant occurrences in Kentucky, 17 in
Tennessee, and 1 in Posey County, Indiana (Table 1). Extant occurrences
in Kentucky are distributed among Clark (1), Franklin (6), and Woodford
(1) Counties, and in Tennessee among Cheatham (5), Davidson (2),
Dickson (1), Jackson (2), Montgomery (3), Smith (1), and Trousdale (2)
Counties. One Tennessee occurrence straddles the county line between
Cheatham and Davidson Counties. There are 19 occurrences in Kentucky
and 10 in Tennessee that have either been extirpated or for which
inadequate information exists to relocate them. Adding the seven
populations that Shea (1993, p. 12-13) treated as either historical or
lacking complete locality information, and which are not represented in
State-maintained databases used to create Tables 1 and 2, these numbers
rise to 20 for Kentucky and 16 for Tennessee. Thus, there is a total of
62 occurrences that have been reported for Short's bladderpod. However,
when reporting percentages of all known occurrences that are now or
historically were in the case of extirpated occurrences, affected by
various threats, we only use the 55 records that have been verified and
are currently tracked in State-maintained databases.
There are 19 extant Short's bladderpod occurrences that are located
on city, State, or federal lands. The Indiana occurrence is on lands
owned by the State of Indiana and managed by the Indiana Department of
Natural Resources (IDNR). A portion of one occurrence in Kentucky is
located in a State nature preserve owned and managed by the Kentucky
State Nature Preserves Commission (KSNPC), and another occurs in a park
owned by the City of Frankfort, where access is limited, but no
specific management is provided for the species or its habitat. In
Tennessee, there are 15 occurrences that are entirely or partially
located on lands owned or leased by the U.S. Army Corps of Engineers
(Corps) adjacent to the Cumberland River. Some of these Corps lands are
wildlife management areas (WMA) cooperatively managed by the Tennessee
Wildlife Resources Agency (TWRA). The plants at EO numbers 29 and 32
are located in TWRA's Cheatham WMA, and those at EO numbers 24 through
27 are located in TWRA's Cordell Hull WMA. Part of one occurrence in
Tennessee is located on lands owned by Ashland City.
Habitat. Short's bladderpod typically grows on steep, rocky, wooded
slopes and talus (sloping mass of rock fragments below a bluff or
ledge) areas. It also occurs along tops, bases, and ledges of bluffs.
The species usually is found in these habitats near rivers or streams
and on south- to west-facing slopes. Most populations are closely
associated with calcareous outcrops (Shea 1993, p. 16). The Short's
bladderpod site in Indiana, where the species is found in a narrow
strip of herbaceous vegetation between a road and forested bank of a
cypress slough (M. Homoya, Natural Heritage Program Botanist, Indiana
Department of Natural Resources (IDNR), December 2012), is unique among
populations of the species. The occurrence in Indiana is within the
Shawnee Hills Section of the Interior Low Plateaus Physiographic
Province (Quarterman and Powell 1978, pp. 30-31), on a site underlain
by undifferentiated outwash from the Wisconsinan glaciation (Indiana
Geologic Survey 2002) as opposed to the calcareous geology on which the
species occurs in Kentucky and Tennessee. The soil at the Indiana site
is Weinbach silt loam, which forms in acid alluvium on river terraces,
and is nearly level with 0 to 2 percent slopes (USDA 1979, p. 89). This
site is on a terrace adjacent to an oxbow swamp formed in an abandoned
meander of the Wabash River (Quarterman and Powell 1978, p. 244).
Kentucky occurrences are located on bluffs and hillsides adjacent
to the Kentucky River or its tributaries within the Bluegrass Section
of the Interior Low Plateaus Province (Fenneman 1938, pp. 411-448;
Quarterman and Powell 1978, pp. 30-31). Extant occurrences in Kentucky
predominantly are found on the Ordovician age Lexington Limestone and
Tanglewood Limestone Members (Kentucky Geological Survey, https://www.arcgis.com/home/item.html?id=d32dc6edbf9245cdbac3fd7e255d3974,
accessed on January 25, 2013), and the Fairmount-Rock outcrop Complex
is the prevalent soil type at most of the sites where the species is
found (U.S. Department of Agriculture (USDA), Soil Survey Geographic
Database, available online at https://soildatamart.nrcs.usda.gov,
accessed on January 30, 2013). Soils of the Fairmount series formed
from
[[Page 47114]]
weathered limestone interbedded with thin layers of calcareous shale
and are shallow, well-drained, and slowly permeable. As implied in the
name of this complex, limestone outcrops are common on the steeply
sloped sites where this soil occurs, especially along river bluffs
(USDA 1985, p. 64).
Tennessee occurrences are located primarily on steep hills or
bluffs adjacent to the Cumberland River within the Highland Rim and
Central (also known as Nashville) Basin Sections of the Interior Low
Plateaus Province (Fenneman 1938, pp. 411-448; Quarterman and Powell
1978, pp. 30-31). Three occurrences in Cheatham County are adjacent to
the Harpeth River near its confluence with the Cumberland River. Extant
occurrences in Tennessee are found across a wider range of geology and
soils than those in Indiana or Kentucky. The Mississippian age Fort
Payne Formation, which includes limestone and calcareous siltstone, and
Warsaw Limestone are the predominant geologic formations underlying
occurrences in Cheatham, Dickson, and Montgomery Counties (Moore et al.
1967, Wilson 1972, Marsh et al. 1973, Finlayson et al. 1980). In
Cheatham and Dickson Counties, the main soil mapped in locations where
Short's bladderpod occurs is simply ``Rock outcrop, very steep'' (USDA,
Soil Survey Geographic Database, available online at https://soildatamart.nrcs.usda.gov, accessed on January 30, 2013). In
Montgomery County, Baxter soils and Rock outcrop and Bodine cherty silt
loam are the soil types on which Short's bladderpod occurs (USDA, Soil
Survey Geographic Database, available online at https://soildatamart.nrcs.usda.gov, accessed on January 30, 2013). Baxter soils
formed from weathered cherty limestone, and where they are mapped as
Baxter soils and Rock outcrop they are steeply sloped and Rock outcrop
can make up as much as 20 percent of the map unit (USDA 1975, pp. 12-
14). Bodine soils are well-drained, cherty soils that formed from
weathered cherty limestone; are steeply sloped; and include areas near
the escarpment adjacent to the Cumberland River floodplain where cherty
limestone bedrock is exposed (USDA 1975, pp. 16-17).
Silurian age limestone and shale of the Waynes Group and the
Brassfield Limestone and Ordovician age limestone of the Leipers and
Catheys Formations are the predominant geologic formations underlying
the occurrences located in Davidson County (Wilson 1979). The dominant
soils on which Short's bladderpod occurs in this county are the Bodine-
Sulphura Complex (USDA, Soil Survey Geographic Database, available
online at https://soildatamart.nrcs.usda.gov, accessed on January 30,
2013), which formed from weathered cherty limestone on sloping to very
steep sites and are somewhat excessively well-drained. Depth to bedrock
within Sulphura soils is less than 16 cm (40 in), but deeper in Bodine
soils, and chert content is high near the surface of these soils (USDA
1981, pp. 46-47).
Ordovician age limestones of the Leipers and Cathey Formations,
Bigby-Cannon Limestone, and Hermitage Formation are the predominant
geologic formations underlying occurrences in Smith, Trousdale, and
Jackson Counties (Wilson et al. 1972, Wilson 1975, Wilson et al. 1980,
Kerrigan and Wilson 2002). In these counties, Short's bladderpod occurs
across a wider range of soil series, all of which are formed from
weathered limestone or interbedded siltstone and limestone on steeply
sloped or hilly sites. The soils are shallow, are rocky, or contain
areas of bedrock outcrop (USDA 2001, pp. 19-20, 28, 59, 64; USDA 2004a,
pp. 22-23, 36-37, 83, 87; USDA 2004b, pp. 21, 75, 82).
Within the physical settings described above, the most vigorous
(Shea 1992, p. 24) and stable (TDEC 2009, p. 1) Short's bladderpod
occurrences are found in forested sites where the canopy has remained
relatively open over time. Common woody species associated with Short's
bladderpod are Acer negundo (box elder), Acer rubrum (red maple),
Aesculus glabra (Ohio buckeye), Celtis laevigata (hackberry), Cercis
canadensis (redbud), Fraxinus Americana (white ash), Juniperus
virginiana (eastern red cedar), Lonicera japonica (Japanese honey
suckle), Parthenocissus quinquefolia (Virginia creeper), Symphoricarpos
orbiculatus (coral berry) and Ulmus americana (American elm). Common
herbaceous associates include Alliaria petiolata (garlic mustard),
Camassia scilloides (wild hyacinth), Chaerophyllum procumbens
(spreading chervil), Delphinium tricorne (dwarf larkspur), Galium
aparine (cleavers), Lamium sp. (dead nettle), Phacelia bipinnatifida
(forest phacelia), Polygonatum biflorum (Solomon's seal), Sedum
pulchellum (stonecrop), Silene virginica (fire-pink), and Verbascum
thapsus (common mullein) (Shea 1993, p. 19).
Biology. Published literature on the biology of Short's bladderpod
is lacking. The species flowers during April and May (Gleason and
Chronquist 1991, p. 187, Shea 1993, p. 20). Dr. Carol Baskin
(Professor, University of Kentucky, pers. comm., December 2012)
observed low fruit set in the Indiana population and, based on lack of
seed production from plants in a greenhouse from which pollinators were
excluded, she concluded that the species likely is self-incompatible.
Self-incompatibility has been reported in other species of Physaria
(Tepedino et al. 2012, p. 142; Edens-Meier et al. 2011, p. 292;
Claerbout et al. 2007, p. 134; Bateman 1955, p. 64), and the molecular
mechanisms underlying self-recognition between pollen and stigma and
subsequent pollen rejection have been well studied in the Brassicaceae
(Takayama and Isogai 2005, pp. 468-474). Dr. Baskin (pers. comm.,
December 2012) also observed that seeds produced by Short's bladderpod
apparently are capable of forming a seed bank, as seeds that were
planted in a greenhouse were observed to germinate and produce
seedlings over several years, rather than all germinating in the year
they were planted.
The pollinators for Short's bladderpod have not been studied, but
Rollins and Shaw (1973, p. 6) reported that bees and flies were
repeatedly observed visiting flowers of other congeners. The majority
of floral foragers observed visiting Physaria filiformis (Missouri
bladderpod) were true bees representing five families, with greater
than 50 percent from the family Halictidae. The families Apidae and
Andrenidae also were well represented among bee pollinators of this
species, the most dependable and frequent of which were ground-nesters.
Several flies of the family Syrphidae also carried Missouri bladderpod
pollen (Edens-Meier et al. 2011, pp. 293). Tepedino et al. (2012, pp.
143-145) found that native ground-nesting bees from the families
Andrenidae and Halictidae were the most reliable pollinators visiting
flowers of three Physaria species, but they reported fewer numbers of
pollen-carrying flies from the families Tachinidae and Conopidae. They
estimated that maximum flight distance ranged from 100 to 1400 meters
(m) (330 to 4593 feet (ft)) for the Andrenids and 40 to 100 m (130 to
330 ft) for the Halictid bees they collected.
Whorled Sunflower
Helianthus verticillatus is a member of the sunflower family known
from Cherokee County, Alabama; Floyd County, Georgia; and McNairy and
Madison Counties, Tennessee. It is a perennial arising from horizontal,
tuberous-thickened roots with slender rhizomes. The stems are slender,
erect, and up to 2 meters (m) (6 feet (ft)) tall. The leaves are
opposite on the lower stem, verticillate (whorled) in groups of
[[Page 47115]]
3 to 4 at the mid-stem, and alternate or opposite in the inflorescence
at the end. Individual leaves are firm in texture and have a prominent
mid-vein, but lack prominent lateral veins found in many members of the
genus. The leaves are linear-lanceolate in shape, narrowing at the tip
to a point, and 7.5 to 18.5 cm (3.0 to 7.2 in.) long and 0.7 to 2.0 cm
(0.3 to 0.8 in.) wide. The flowers are arranged in a branched
inflorescence typically consisting of 3 to 7 heads. The heads are about
1 cm high (0.4 in.), are about 1.5 cm (0.6 in.) wide, and have deep
yellow ray flowers and lighter yellow disk flowers. The seeds are 0.4
to 0.5 cm (0.16 to 0.2 in.) long.
Several members of the aster family are similar in appearance to
whorled sunflower, with minor morphological differences being apparent.
Helianthus grosseserratus is similar to whorled sunflower but its
leaves typically are arranged in an alternating pattern, which differs
from the whorled arrangement of H. verticillatus. Helianthus
angustifolius can be confused with H. verticillatus but it has narrower
leaves and reddish disk flowers, as opposed to the yellow disk flowers
of H. verticillatus (Schotz 2001, p. 1). Helianthus giganteus often
exhibits whorled leaves, but H. verticillatus leaves have only the
midvein prominent while H. giganteus has lateral veins evident on the
leaves (Matthews et al. 2002, p. 22).
Taxonomy. Whorled sunflower was described by J.K. Small (1898, p.
479), based on a collection by S.M. Bain from Chester County,
Tennessee, in 1892. Small distinguished it from the related H.
giganteus by its smooth and hairless stems; narrow, entire leaf blades;
and narrowly linear-lanceolate involucre (a collection or rosette of
bracts subtending a flower cluster, umbel, or the like) bracts (a
leaflike or scalelike plant part, usually small, sometimes showy or
brightly colored, and located just below a flower, a flower stalk, or
an inflorescence). No additional collections of this species had been
made when Beatley (1963, p. 153) speculated that the specimens (which
lacked basal parts and mature seeds) from this single collection site
perhaps represented a single aberrant individual formed from
hybridization of an opposite- and alternate-leaved Helianthus species.
With no new material to examine, Heiser et al. (1969, p. 209) and
Cronquist (1980, p. 36) accepted Beatley's suggestion that whorled
sunflower was a hybrid.
The rediscovery of the species in 1994, in Georgia, provided ample
material for reexamination of this species' taxonomic status. Plants
throughout these new populations were found to conform to the
morphology of the type collection of whorled sunflower. Morphological
studies and root-tip chromosome counts by Matthews et al. (2002, pp.
17-23) validated this taxon's status as a distinct, diploid species.
The taxonomic validity of this species was also confirmed through
genetic studies by Ellis et al. (2006, pp. 2345-2355). Their studies
showed through comparative genetic studies with its putative parents,
H. grosseserratus and H. angustifolius, that whorled sunflower is a
good taxonomic species of non-hybrid origin (Ellis et al. 2006, pp.
2351-2352).
Distribution and Status. There are four whorled sunflower
populations known to be extant, each consisting of multiple tracked
subpopulations (Table 3) (Alabama Natural Heritage Program (ANHP) 2012,
Georgia Department of Natural Resources (GDNR), TNHID 2012). In Floyd
County, Georgia, there is one population comprised of four
subpopulations. There is one population in Cherokee County, Alabama,
comprised of two subpopulations. Populations in Georgia and Alabama are
less than 2 km (1.2 mi) apart. In Tennessee, there is one population
comprised of six subpopulations in McNairy County and one population
comprised of four subpopulations in Madison County. Table 3 lists these
populations and subpopulations, and relates them to EO numbers used by
State conservation agencies to track their status. The population in
Floyd County, Georgia, is located on lands owned by The Campbell Group,
a timber investment management organization. This site is referred to
as the Coosa Valley Prairie and is protected by a conservation easement
held by The Nature Conservancy, which jointly manages the property with
The Campbell Group. All other sites also are on private lands but are
not protected.
Table 3--List of Whorled Sunflower Populations and Subpopulations by State and County, With Corresponding Site
Names and Element Occurrence (EO) Numbers From State Conservation Agency Databases in Alabama, Georgia, and
Tennessee
----------------------------------------------------------------------------------------------------------------
Subpopulation
Population (County, State) number(s) Site name Heritage EO Number
----------------------------------------------------------------------------------------------------------------
Cherokee, AL............................ 1 Kanady Creek Prairie....... AL--1
2 Locust Branch Prairie...... AL--2
Floyd, GA............................... 1 Jefferson Road Wet Prairie. GA--1
2 Kanady Creek Wet Prairie... GA--4
3 Upper Mud Creek Wet GA--5
Prairies.
4 Sunnybell Prairie.......... GA--7
Madison, TN............................. 1-6 Turk Creek................. TN--2
McNairy, TN............................. 1-4 Prairie Branch............. TN--3
----------------------------------------------------------------------------------------------------------------
Status surveys have been conducted for this species throughout its
range (Nordman 1998, pp. 1-17; 1999, pp. 1-5; Schotz 2001, pp. 1-14;
Allison 2002, pp. 1-2; Lincicome 2003, pp. 1-2). Despite these
extensive surveys, the number of known populations remains low. Schotz
(2001, pp. 1, 10) located 1 new population out of 44 attempts,
representing a success rate of only 2 percent. Surveys during 2000 and
2002 in Tennessee were unsuccessful at locating any additional sites
(Lincicome 2003, pp. 1-2). Surveys in 2006 resulted in discovery of the
population in McNairy County, Tennessee (Tennessee Division of Natural
Areas 2008, p. 2).
Initial efforts to estimate population sizes of whorled sunflower
relied on counting individual stems (Allison 2002, pp. 3-8; Schotz
2001, pp. 8-10); however, due to the species' clonal growth habit, stem
counts overestimate the true number of genetically distinct individuals
(genets). Ellis et al. (2006, p. 2349) found that the genetic
population size is much smaller than the number of stems in a
population and that a more accurate population census could be made at
most whorled sunflower sites
[[Page 47116]]
by counting obvious clusters of stems rather than individual stems.
However, Mandel (2010, p. 2056) reported that individual clusters were
much less distinct in a portion of the Alabama site she sampled.
Ellis et al. (2006, p. 2351) counted 70 distinct clusters at the
site in Madison, Tennessee, which closely equated to 70 separate
individuals through genetic analyses; however, not all clusters were
sampled at this site (Mandel, pers. comm., 2012). At the McNairy
County, Tennessee, population, 36 clusters of plants were found growing
along creek banks at the unplowed edges of cultivated crop fields and
extending into a railroad right-of-way (Tennessee Division of Natural
Areas 2008, p. 3). Mandel (2010, p. 2056) sampled 19 clusters at the
McNairy County population and determined these represented 24 genets;
however, only two of the four subpopulations mapped at this population
were sampled (Mandel, pers. comm., 2012).
Mandel (2010, p. 2058) sampled the Alabama subpopulation number 1
(Table 3) using two methods. In one portion of the site, leaf tissue
was collected from 15 distinct clusters, which represented 24 genets.
However, because distinct clusters were not obvious in another portion
of this subpopulation, Mandel (2010, p. 2058) sampled leaves from the
first 100 stalks encountered in a 1-meter-wide transect run through the
largest patch of whorled sunflower in that area. These 100 stalks were
within an approximately 11-m (40-ft) long portion of this transect, and
represented 46 distinct genets. Mandel (2010, p. 58) estimated that 400
stalks were present in this area and that the total number of genets
was between 100 and 200. However, more recently only 79 stems,
distributed among 8 clusters, were found at this site (Alabama Natural
Heritage Program 2011, p. 11).
Mandel (2010, p. 2056) sampled 15 clusters growing in a ``wet
prairie'' at the Georgia site, presumably representing EO number 1 from
the Georgia Natural Heritage Program database (Table 3). It was
determined that these clusters represented 18 genets (Mandel 2010, p.
2058), but apparently the other three subpopulations present at this
population were not sampled. The true number of genets at this site is
likely much greater, as others have reported vigorous growth of whorled
sunflower in response to prescribed fires that are used to manage the
Coosa Valley Prairie conservation easement area (M. Hodges, Georgia
Director of Stewardship, The Nature Conservancy, pers. comm. May 2012;
T. Patrick, Botanist, Georgia Department of Natural Resources, pers.
comm. February 2012).
Based on the work of Ellis (2006) and Mandel (2010), summarized
above, at one time Alabama supported the largest population with an
estimated 100 individuals at the Kanady Creek Prairie site, where
whorled sunflower was first found to occur in the State. However,
Schotz (2011, p. 11) found only 79 stems, distributed among 8 clusters,
at this site in 2011. Mandel (2010) sampled only portions of the
Georgia and Tennessee populations, thus underestimating their sizes.
Whorled sunflower likely is now most abundant in Georgia due to
population growth in response to habitat management by The Nature
Conservancy and The Campbell Group at the Coosa Valley Prairie. Schotz
estimated approximately 175 to 200 stems were present at the second
Alabama site in September 2008 (Schotz pers. comm. 2009), but there
were only 42 stems found at this site in 2011 (Schotz 2011, p. 14). No
estimate of individual plants is available for this site.
Habitat. Whorled sunflower is found in moist, prairie-like
remnants, which in a more natural condition exist as openings in
woodlands and adjacent to creeks. Today, the only whorled sunflower
site where these habitat conditions are present over a relatively large
area is located in the Coosa Valley Prairie of northwest Georgia, where
the species occurs in prairie openings and woodlands interspersed among
lands managed for pulpwood and timber production. At one of the Alabama
subpopulations, whorled sunflower occurs in a narrow, open strip of
vegetation between a roadside and adjacent forest. The second Alabama
subpopulation occurs along a small intermittent stream and adjacent
floodplain, in a site where an immature hardwood forest was harvested
in 1998. Whorled sunflower and associated prairie species responded
favorably to the timber removal, but the site was soon converted into a
loblolly pine plantation and the planted seedlings have grown into a
young, dense stand into which little light penetrates. As of 2012,
there were few whorled sunflower plants or prairie associates present
at this site. Known populations of this species in Tennessee are
relegated mostly to narrow bands of habitat between cultivated fields
and creeks and adjacent to roads and railroad rights-of-way. The
largest concentration of plants in Tennessee is found at the Madison
County population, in a 1-ha (2.5-ac) patch of remnant, wet prairie
habitat wedged between US Highway 45 and a railroad right-of-way.
The Alabama and Georgia populations are located on flat to gently
rolling uplands and along stream terraces in the headwaters of Mud
Creek, a tributary to the Coosa River. In Tennessee, the Madison County
population occurs along Turk Creek, a tributary to the South Fork
Forked Deer River, and in adjacent uplands. The McNairy County
population occurs along Prairie Branch, a headwater tributary to Muddy
Creek in the Tuscumbia River drainage.
We used the Natural Resources Conservation Service's Web Soil
Survey to determine the soil types on which whorled sunflower
populations occur across its range (USDA, Web Soil Survey, available
online at https://websoilsurvey.nrcs.usda.gov/app/HomePage.htm, accessed
on January 30, 2013). The most prevalent soils where the species occurs
in Georgia are Conasauga, Lyerly, Townley, and Wolftever silt loams and
Dowellton silty clay loam. The silt loam soils all formed from
weathered limestone or shale, and occupy various land forms from broad
upland ridges to low stream terraces. These soils share the
characteristics of being moderately well-drained to well-drained, being
slightly to extremely acid, and having low to moderate fertility and
organic matter content and clayey subsoils (USDA 1978a, pp. 24-54). The
Dowellton silty clay loam formed in alluvium (soil material deposited
by running water) on low stream terraces and upland depressions is
poorly drained, is moderate in fertility and organic content, is
neutral to strongly acid, and has a clayey subsoil (USDA 1978a, pp. 28-
29).
Alabama subpopulations inhabit the Gaylesville silty clay loam, a
deep, poorly drained, slowly permeable soil formed from limestone on
floodplains and depressed areas in limestone valleys (USDA 1978b, p.
20). These soils are strongly to extremely acid, with low natural
fertility and medium organic content (USDA 1978b, p. 20). Conasauga
silt loams, discussed above, lay upslope of the Gaylesville soils at
the Alabama whorled sunflower sites.
In Madison County, Tennessee, the population is primarily found on
Falaya silt loam, which are poorly drained soils that formed in
alluvium derived from loess (loamy soil material believed to be
deposited by wind) and are strongly to very strongly acid (USDA 1978,
p. 44). The McNairy County, Tennessee, population occurs on Iuka and
Enville fine sandy loam soils, both of which occupy floodplains and are
occasionally flooded during winter and early spring (USDA 1997, pp. 73-
76).
The list of associated species in these habitats indicates a
community with
[[Page 47117]]
strong prairie affinities. Dominant grasses of the tall grass prairie
are present including Schizachyrium scoparium (little bluestem),
Sorghastrum nutans (Indian grass), Andropogon gerardii (big bluestem),
and Panicum virgatum (switch grass). Other common herbaceous associates
include Bidens bipinnata (Spanish needles), Carex cherokeensis
(Cherokee sedge), Hypericum sphaerocarpum (roundseed St. Johnswort),
Helianthus angustifolius (swamp sunflower), Helenium autumnale (common
sneezeweed), Lobelia cardinalis (cardinal flower), Pycnanthemum
virginianum (Virginia mountain mint), Physostegia virginiana (obedient
plant), Saccharum giganteum (sugarcane plumegrass), Silphium
terebinthinaceum (prairie rosinweed), Sporobolus heterolepis (prairie
dropseed), and Symphyotrichum novae-angliae (New England aster)
(Tennessee Division of Natural Areas 2008, p. 5; Matthews et al. 2002,
p. 23; Schotz 2001, p. 3). Some of these areas are also habitat for a
number of other rare species including Marshallia mohrii (Mohr's
Barbara's buttons), which is federally listed as threatened.
Biology. There is little published information available concerning
the biology of the whorled sunflower, and the cause for its current
rarity is not known. Ellis et al. (2006, pp. 2349-2350) investigated
genetic diversity in the Georgia, Alabama, and Madison County,
Tennessee, populations of whorled sunflower and found high levels of
genetic diversity at the population and species levels despite its
apparent rarity. They speculated that this is indicative of a species
that was more widespread in the past and perhaps became rare relatively
recently (Ellis et al. 2006, pp. 2351-2352). Whorled sunflower
populations exhibited moderate levels of differentiation based on
markers that are presumed to be selectively neutral, and since these
populations are geographically distinct and ecological conditions vary
somewhat among them Ellis et al. (2006, p. 2353) concluded that they
likely are as differentiated, if not more so, at adaptive loci (the
specific location of a gene or DNA sequence on a chromosome).
Whorled sunflower is a self-incompatible, clonal perennial and
flowers from August into October (Matthews et al. 2002, pp. 17-20;
Ellis and McCauley 2008, p. 1837). The species is easily cultivated and
seed germination is high in the laboratory. Upon transplanting, this
species has been shown to reproduce rapidly from rhizomes (a horizontal
underground stem that produces roots and shoots), creating dense
colonies. The stems can reach over 4 m (13 ft) in height (Matthews et
al. 2002, pp. 17-20).
Ellis and McCauley (2008, p. 1837) investigated whether there were
differences among populations of whorled sunflower with respect to
achene viability and germination rates and whether those differences
might have a genetic basis. They conducted this experiment for two
generations of plants, the second generation produced from intra-
population crosses of first generation plants. They also explored
whether isolation of populations from one another could have fitness
consequences, by conducting inter-population crosses and evaluating
whether they found: (1) Evidence of genetic rescue expressed as higher
fitness of hybrid individuals as compared to any or all of the parental
populations; and (2) evidence of outbreeding depression. Their study
included material from the Alabama, Georgia, and Madison County,
Tennessee, populations. However, they were unsuccessful in cultivating
plants from the Georgia population, where the flower heads contained
few viable achenes, which produced low germination rates (Ellis and
McCauley 2008, pp. 1837-1838).
The number of crosses that produced no viable achenes was higher in
the intra-population Tennessee crosses than in any other pair of
crossings. Those achenes that were produced by first generation
Tennessee intra-population crosses exhibited lower germination rates
than Alabama achenes, and second generation Tennessee achenes from
intra-population crosses exhibited both lower viability and germination
rates than the Alabama achenes. However, survival rates of germinated
achenes did not differ among these populations in either generation
(Ellis and McCauley 2008, p. 1840). Ellis and McCauley (2008, p. 1840)
suggested three possible mechanisms that could explain these results,
none of which are mutually exclusive: (1) Limited mate availability in
the Tennessee population due to limited diversity of self-
incompatibility alleles; (2) more extensive inbreeding within the
Tennessee population; or (3) differential adaptation between the two
populations.
When Tennessee plants were crossed with pollen from Alabama plants,
the second generation mean achene viability and germination rates were
equal to or greater than those of Alabama intra-population crosses or
Alabama plants crossed with pollen from Tennessee plants. Mean achene
viability of Tennessee intra-population second generation crosses was
lower than all other groups and germination rates were lower than both
Alabama intra-population crosses and Alabama plants crossed with pollen
from Tennessee plants (Ellis and McCauley 2008, pp. 1839-1840).
Based on their results, Ellis and McCauley (2008, p. 1841)
concluded that populations of whorled sunflower are not interchangeable
with respect to phenotypic fitness-related characters (i.e., achene
viability and germination rates) and suggested that the potential
exists for genetic rescue of the Tennessee population by transplanting
either seeds or seedlings produced from crosses between Tennessee and
Alabama plants into the Tennessee population.
Fleshy-fruit Gladecress
Leavenworthia crassa is a glabrous (morphological feature is
smooth, glossy, having no trichomes (bristles or hair-like structures))
winter annual known from Lawrence and Morgan Counties, Alabama. It
usually grows from 10 to 30 cm (4 to 12 in) tall. The leaves are mostly
basal, forming a rosette, and entire to very deeply, pinnately
(multiple leaflets attached in rows along a central stem) lobed or
divided, to 8 cm (3.1 in) long. Flowers are on elongating stems, and
the petals are approximately 0.8 to 1.5 cm (0.3 to 0.6 in.) long,
obovate to spatulate, and emarginate (notched at the tip). Flower color
is either yellow with orange or white with yellow, usually with both
color forms intermixed in a single population. The fruit is globe-
shaped or slightly more elongate and about 1.2 cm (0.5 in) long with a
slender beak at the tip, which is 0.25 to 0.60 cm (0.1 to 0.24 in) in
length. Seeds are dark brown, nearly round in shape and winged.
Taxonomy. Fleshy-fruit gladecress was described by Rollins in 1963,
from material collected in 1959, from Morgan County, Alabama. Rollins
(1963, pp. 61-68) delineated the species into two varieties (var.
crassa and var. elongata) based on differences in fruit length.
However, herbarium and field studies have shown var. elongata to have
variation in fruit length within the range of fruit lengths for var.
crassa (McDaniel and Lyons 1987, p. 2-3). Thus, the species is treated
as one taxon throughout this document. This taxon was brought to the
attention of the scientific community in 1957, by venerable botanist
Reed C. Rollins, who distinguished the taxon from similar species based
on reproductive morphology.
Fleshy-fruit gladecress's globular to oblong fruit with a smooth
exterior distinguishes it from another gladecress
[[Page 47118]]
species, Leavenworthia alabamica (Alabama gladecress), which has a much
more elongated linear fruit with corrugated surfaces. Alabama
gladecress also does not usually have the yellow and orange flower
forms found mixed in populations of fleshy-fruit gladecress (McDaniel
and Lyons 1987, p. 10).
Distribution and Status. Fleshy-fruit gladecress is endemic to a
21-km (13-mi) radius area in north central Alabama in Lawrence and
Morgan Counties (Rollins 1963, p. 63). A 1961 record from Lauderdale
County has never been confirmed (McDaniel and Lyons 1987, p. 6).
Surveys by Lyons (in litt. 1981 to R. Sutter), McDaniel and Lyons
(1987, p. 5-6), and Hilton (1997, p. 12) were unsuccessful at locating
a number of historical sites for fleshy-fruit gladecress. McDaniel and
Lyons (1987) failed to locate eight sites previously reported by
Rollins (1963, p. 63), and Lloyd (1965) and Hilton (1997, p.12) were
unsuccessful at locating seven sites listed in McDaniel and Lyons
(1987, p. 5-6).
Currently there are six known extant occurrences of fleshy-fruit
gladecress documented, three each in Morgan and Lawrence Counties,
Alabama (Table 4). One of these occurs on U.S. Forest Service (USFS)
lands, where it is formally protected. The majority of other sites are
actively grazed, a practice that has, for the most part, maintained
favorable growing conditions for the species. However, adjusting
grazing patterns to take place during the species' dormant cycle would
greatly reduce potential mortality of reproducing plants while
maintaining ideal habitat conditions.
Table 4--Location, Site Names and Descriptions, and Element Occurrence (EO) Ranks for Known Extant Fleshy-Fruit
Gladecress Occurrences
----------------------------------------------------------------------------------------------------------------
Population Historic site
County designation EO Rank description Land ownership
----------------------------------------------------------------------------------------------------------------
Lawrence........................ Bluebird Glades..... D......... Described by ALNHP in Private & State
1995 as approx. 0.2-ha ROW.
(0.5-ac) site with
1200 plants; by 2009
was reduced to 600
plants.
Stover Branch Glades C......... Two subpopulations, Private.
most in pasture, 3.16
ha (7.8 ac); 2,200 to
2,500 plants;
maintained by
livestock management,
found in 1961.
Indian Tomb Hollow A......... 0.46-ha (1.1-ac) site Federal--USFS.
Glade. with 1,200 to 1,300
plants; discovered
1977.
Morgan.......................... Cedar Plains South.. C......... 0.04-ha (0.1-ac) site Private.
with 75 to 100 plants;
discovered 1968.
Cedar Plains North.. B......... 1.7-ha (4.2-ac) site Private.
with 5,000 to 6,000
plants; discovered
1968.
Massey Glade........ C......... 2.75-ha (6.8-ac) site Private.
with 2,300 to 2,500
plants; discovered
1961.
----------------------------------------------------------------------------------------------------------------
ALNHP is the Alabama Natural Heritage Program.
ROW is right-of-way.
The Alabama Natural Heritage Program determines EO ranks ranging
from A to D for sites and populations of rare species, with A
indicating the status of the EO is considered to be excellent, B good,
C marginal, and D poor. The EO rank is based on a combination of
standardized criteria including quality, condition, viability, and
defensibility. Hilton (1997, pp. 13-26) developed the specific criteria
for determining EO ranks for fleshy-fruit gladecress and its habitat.
Based on these criteria, only one of the six occurrences is A-ranked.
It consists of an estimated 1200+ plants in a relatively undisturbed
glade (Schotz 2009, p. 10). Of the remaining occurrences, one has
approximately 5,000 to 6,000 plants, but is B-ranked because the site
where it is located is heavily grazed. Three occurrences are C-ranked
(2 occurrences have approximately 2400 plants in a degraded glade
community; the other occurrence has 75 to 100 plants but is located in
high-quality habitat), and one is D-ranked (600 plants in a residential
area with no potential for habitat restoration) (Schotz 2009).
Habitat. This species is a component of glade flora and occurs in
association with limestone outcroppings. The terms ``glade'' and
``cedar glades'' are used interchangeably to refer to shallow-soiled,
open areas that are dominated by herbaceous plants and characterized by
exposed sheets of limestone or gravel. Eastern red cedar (Juniperus
virginiana) trees are frequent in the deeper soils along the edges of
the glades (Hilton 1997, p. 1; Baskin et al. 1986, p. 138; Baskin and
Baskin 1985, p. 1). Glades can vary in size from as small as a few
square meters to larger than 1 square kilometer (km\2\) (0.37 square
miles (mi\2\)) and are characterized as having an open, sunny aspect
(lacking canopy) (Quarterman 1950, p. 1; Rollins 1963, p. 5).
Historically, glades in northern Alabama occurred as glade complexes
where sparsely vegetated patches of exposed, or nearly exposed,
limestone occurred in a matrix of woody vegetation to form a mosaic of
habitats grading into one another (Hilton 1997, pp. 1, 5, 64).
Herbaceous diversity was irregular over these complexes, affected by
changes in soil gradient and moisture, and the presence or absence of a
woody vegetation component. Few undisturbed examples of this community
type remain (Hilton 1997, pp. 5, 8; McDaniel and Lyons 1987, p. 11;
Baskin and Baskin 1985, p. 1; Rollins 1963, p. 5-6).
Populations of fleshy-fruit gladecress are now located in glade-
like remnants exhibiting various degrees of disturbance, including
pastures, roadside rights-of-way, and cultivated or plowed fields
(Hilton 1997, p. 5). As with most of the cedar glade endemics, fleshy-
fruit gladecress exhibits weedy tendencies, and it is not uncommon to
find the species growing in altered habitats. However, none of the
cedar glade endemics appear to have spread very far from their original
glade habitats; thus the geographic range of fleshy fruit gladecress is
probably very similar to what it was in pre-settlement times (Baskin et
al. 1986, p. 140).
All species within the small genus Leavenworthia are adapted to the
unique physical characteristics of glade habitats, perhaps the most
important of these being a combination of shallow
[[Page 47119]]
depth and high calcium content of soils and their tendency to have
temporarily high moisture content at or very near the surface (Rollins
1963, pp. 4-6). Typically, only a few inches of soil overlie the
bedrock, or, in spots, the soil may be almost lacking and the surface
barren. The glade habitats that support all Leavenworthia species are
extremely wet during the late winter and early spring, and become
extremely dry in summer (Rollins 1963, p. 5).
In northern Alabama, cedar glades primarily are distributed within
the Moulton Valley subdivision of the Interior Low Plateau
Physiographic Province, and a few glades are scattered up the Eastern
Valley subdivision of the Tennessee Valley (Hilton 1997, p. 1). Most of
these glades are concentrated in the Moulton Valley, a level area
underlain by Mississippian age limestone stretching across Morgan,
Lawrence, Franklin, and Colbert Counties in northwestern Alabama.
Glades occur in association with outcrops of Bangor Limestone and
typically are level with exposed sheets of limestone or limestone
gravel interspersed with fingers of cedar-hardwood vegetation. The
Bangor Limestone underlying the Moulton Valley tapers to an end in
eastern Morgan County, where it meets the sandstone of Brindley
Mountain. Limestone is often near the soil surface, and can be seen in
rocky cultivated fields and as small outcroppings at the base of low-
lying forested hills (Hilton 1997).
Biology. Fleshy-fruit gladecress is an annual, spring-flowering
member of the mustard family (Brassicaceae). As an annual, the seeds
germinate in the fall, overwinter as rosettes, and commence a month-
long flowering period beginning in mid-March. The first seeds mature in
late April, and during most years the plants dry and drop all of their
seeds by the end of May. It is unlikely that all seeds produced in
spring germinate the next fall, but the length of dormancy in the soil
is not known (McDaniel and Lyons 1987, p. 10); thus we do not know
whether the species is capable of forming a seed bank. Native bees in
the families' Andrenidae and Halictidae (sweat bees), including the
species Halictus ligatus (sweat bee), were observed carrying pollen
from Leavenworthia crassa (fleshy-fruit gladecress) and L. alabamica
(Alabama gladecress) in northern Alabama (Lloyd 1965).
Summary of Factors Affecting the Species
Section 4 of the Act (16 U.S.C. 1533), and its implementing
regulations at 50 CFR part 424, set forth the procedures for adding
species to the Federal Lists of Endangered and Threatened Wildlife and
Plants. Under section 4(a)(1) of the Act, we may list a species based
on any of the following five factors: (A) The present or threatened
destruction, modification, or curtailment of its habitat or range; (B)
overutilization for commercial, recreational, scientific, or
educational purposes; (C) disease or predation; (D) the inadequacy of
existing regulatory mechanisms; and (E) other natural or manmade
factors affecting its continued existence. Listing actions may be
warranted based on any of the above threat factors, singly or in
combination. Each of these factors is discussed below.
Short's Bladderpod
A. The Present or Threatened Destruction, Modification, or Curtailment
of Its Habitat or Range
Shea (1993, pp. 22-23 and 42-92) and Tennessee Department of
Environment and Conservation (2009, p. 1-3) discussed several threats
that have destroyed or modified Short's bladderpod habitat and could
cause further habitat loss or modification in the future. These include
transportation right-of-way construction and maintenance; impoundments
and reservoir water level manipulation; overstory shading due to forest
succession; competition and shading from invasive, nonnative plant
species; trash dumping; commercial and residential construction; and
livestock grazing. Predictions of increased frequency, duration, and
intensity of droughts across the species' range, and increased flooding
in the Midwest region, could portend adverse effects for Short's
bladderpod and its habitat. We discuss each of these threats in greater
detail below.
Transportation Right-of-Way Construction and Maintenance
During the status survey for this species, Shea (1993, p. 22)
observed that Short's bladderpod habitat at three sites (Kentucky EO 7;
Tennessee EOs 7, 14) had been destroyed or degraded by road
construction or maintenance activities. Neither of these Tennessee
occurrences is extant today (TNHID 2012). Shea (1993, p. 60) observed
48 plants at Kentucky EO 7 in 1992, but noted that the population had
been much more extensive prior to improvements of U.S. 421. Shea (1993,
p. 22) also indicated that roadside maintenance posed a continuing
threat to the species at this location. Although approximately 100
Short's bladderpod plants were observed on a steep slope above the road
cut adjacent to Kentucky EO 7 in 2004 (KNHP 2012), no plants were found
at the base of the bluff, where 21 plants had been observed in 1992
(Shea 1993, p. 60) before the road cut had altered the habitat. Poorly
timed mowing or indiscriminate herbicide application along the road cut
at the base of this bluff could cause mortality of seedlings produced
there from seeds that are dispersed from the plants on the slope above.
According to data from the KNHP (2012), a road cut was present in 2004,
and no Short's bladderpod could be found at Kentucky EO 2, where in
1992 Shea (1993, p. 52) observed 11 Short's bladderpod plants and
observed no apparent threats to the population. Much of the habitat
downslope of a road, where Tennessee EO 20 once occurred but is no
longer extant, was found to be covered with rip rap in 2008, and the
remaining habitat above and below the road was overgrown (TDEC 2009, p.
10). Road construction destroyed suitable habitat around Tennessee EO
23, and Short's bladderpod is no longer present at the site (TNHID
2012). Based on these data, five Short's bladderpod occurrences (9
percent) have been lost to habitat destruction or modification
associated with road construction or maintenance.
Shea (1993, p. 22) identified roadside maintenance as a threat to
12 occurrences, including two discussed above: Indiana EO 1; Kentucky
EOs 1 through 4, 7, 19, and 23; and Tennessee EOs 2, 4, 10, and 22. In
addition, Kentucky EO 27 is located along a mowed roadside (KNHP 2012),
and TDEC (2009, p. 2) reported that Tennessee EOs 3 and 15 could be
affected by roadside maintenance. Indiana EO 1 is an extant roadside
occurrence, where the species' persistence depends on periodic clearing
of competing vegetation and associated soil disturbance to prevent
succession of the vegetation at the site to a forested condition that
would be unsuitable for Short's bladderpod (Homoya, pers. comm.,
December 2012). Nonetheless, poorly timed mowing or indiscriminate
herbicide application could negatively affect this occurrence by
disrupting reproductive cycles or causing direct mortality of Short's
bladderpod plants. In total, roadside maintenance has been identified
as a threat to 15 occurrences.
Short's bladderpod is considered extirpated from four of the eight
sites in Kentucky where roadside maintenance has been identified as a
threat to the species. Neither Kentucky EO 2, lost to road construction
as discussed above, nor EO 3 is extant. No plants were
[[Page 47120]]
found at Kentucky EO 3 during searches in 2004 and 2008; however, only
a few plants had been observed here in 1994 and earlier (KNHP 2012),
and the cause for the species' current absence is not known. Despite
the presence of 17 Short's bladderpod plants at Kentucky EO 19 during
2005, none were found during visits in 2004 and 2011 (KNHP 2012). While
roadside maintenance could have contributed to loss of this population,
observations by Kentucky Natural Heritage Program (2012) indicate that
shading or competition from invasive species is likely a primary cause.
Short's bladderpod was last seen at Kentucky EO 27 in 1993, when seven
plants were found along a mowed roadside dominated by fescue and other
weeds (KNHP 2012). This occurrence was determined to be extirpated
during a 2011 site visit by KNHP (2012) staff.
Short's bladderpod remains extant at four of the eight sites in
Kentucky where roadside maintenance has been identified as a threat to
the species. Kentucky EO 1 is considered extant, but only three Short's
bladderpod plants--two in 1992, and one in 2009--have been observed at
this site since the species was first observed there in 1975. Kentucky
EO 4 was treated as two separate populations by Shea (1993, pp. 62-65),
which are now tracked as a single occurrence (KNHP 2012). While some
plants at the base of the cliff where Kentucky EO 4 is located are
vulnerable to roadside mowing or herbicide application, many of the
plants are on the cliff face and associated ledges, and no impacts from
roadside maintenance have been documented. Short's bladderpod abundance
at this occurrence has ranged from a low of approximately 56
individuals in 1998, to a high of at least 400 individuals in 2004
(KNHP 2012). As discussed above, there were approximately 100 plants
observed above the road cut at Kentucky EO 7, but roadside maintenance
could prevent plants from becoming established at the base of the road
cut. Kentucky EO 23 has ranged in abundance from a low of 60 plants in
2008, to a high of at least 430 plants in 2001. In 2011, there were
more than 500 seedlings present at this site, but no flowering plants
were observed. While this occurrence is located near a roadside, there
have been no documented impacts from roadside maintenance.
Short's bladderpod is considered extirpated from two of the seven
sites in Tennessee where roadside maintenance has been identified as a
threat to the species. At Tennessee EO 2, TDEC (2009, p. 5) found the
habitat to be too overgrown and Short's bladderpod absent during a
search in 1998, and no plants were found during a monitoring visit in
2008. As noted above, Short's bladderpod was no longer present when
TDEC (2009, p. 10) observed in 2008 that the roadside habitat at
Tennessee EO 20 had been covered with rip rap and the remaining habitat
above and below the road was overgrown.
Short's bladderpod remains extant at five of the seven sites in
Tennessee where roadside maintenance has been identified as a threat to
the species. More than 500 Short's bladderpod plants were found at
Tennessee EO 3 in 2008 (TDEC 2009, p. 6), where Shea (1993, p. 89)
found 40 plants in 1992. This occurrence is located along a south-
facing wooded slope, north of the Cumberland River, but very little of
its habitat would be vulnerable to maintenance associated with the road
right-of-way to the immediate west. Tennessee EOs 4 and 10 are located
along a roadside approximately 0.5 km (0.3 mi) apart, and both
occurrences are estimated to number in the hundreds to thousands of
plants (TDEC 2009, p. 6-8). While roadside maintenance could adversely
affect plants located along the base of the roadside bluffs on which
they occur, the majorities of these occurrences are located on ledges
and bluff tops where roadside maintenance would be unlikely to affect
them. Tennessee EO 15 is a small occurrence located adjacent to a
bridge, on a steep limestone bluff overlooking the Harpeth River. While
no impacts from roadside maintenance have been observed, no more than
20 plants have ever been counted at this occurrence. Biologists from
TDEC (2009, p. 11) found approximately 35 plants at Tennessee EO 22,
where Shea (1993, p. 85) found 43 reproductive plants in 1992. No
impacts from roadside maintenance were noted during this site visit.
Four Short's bladderpod occurrences (7 percent) apparently have
been lost to road construction or roadside maintenance. While 10 of the
known extant occurrences (38 percent) are located along roadsides,
where maintenance activities such as mowing or herbicide application
could affect them, there have been few documented examples of such
effects. In many roadside locations, Short's bladderpod occurs on steep
slopes or bluffs, where roadside maintenance would be unlikely to
affect the species unless the road was widened, requiring alteration or
removal of the slope or bluff. Moreover, well-timed and carefully
executed right-of-way maintenance intended to control vegetation
encroachment could be beneficial by reducing shading and competition.
Nonetheless, the potential exists for road widening projects or
vegetation management efforts along road rights-of-way to destroy or
modify habitat, cause mortality of individual plants, or diminish
reproductive output at a large proportion of sites where the species
occurs.
There are seven extant Short's bladderpod occurrences, and three
sites from which the species is thought to be extirpated, located in or
adjacent to the Old Tennessee Central Railroad right-of-way (TDEC 2009,
p. 3, TNHID 2012), portions of which are not actively used or
maintained or have been sold to other rail companies. There were
hundreds to thousands of Short's bladderpod plants each at three of
these occurrences (Tennessee EOs 1, 10, and 17) when TDEC (2009, p. 4)
monitored the species in 2008. The Nashville Area Metropolitan Planning
Organization (NAMPO) (2010, p. 98) 2035 Regional Transportation Plan
reported that the Old Tennessee Central Railroad, which follows the
Cumberland River and passes through Ashland City, was found to be the
most practical alignment for a proposed commuter rail to improve
intercity commute options between the cities of Nashville and
Clarksville, Tennessee. While no plans have been produced for
developing this proposed commuter rail system, the 2035 Regional
Transportation Plan states that this transportation option should be
developed by 2017 (NAMPO 2010, p. 98). Habitat modification or
destruction resulting from such development could potentially affect 27
percent of the known extant occurrences of the species, including some
occurrences where the species is most abundant.
Flooding and Water Level Fluctuation
Shea (1993, pp. 22-23) and TDEC (2009, p. 2) noted that
impoundments and artificial water level manipulation threatened several
Short's bladderpod occurrences. This threat might be better
characterized as flooding and water level fluctuation, regardless of
cause, as some occurrences in free-flowing river reaches are vulnerable
to this threat. For example, the Indiana occurrence is located near an
oxbow lake that was created in a relict channel of the Wabash River,
and it is periodically inundated by floodwaters from the river. In
2011, this occurrence was subjected to a prolonged flood that killed
most of the Short's bladderpod plants at this location (Homoya, pers.
comm., November 2012). There were thousands of seedlings present at
this site in 2010, and this flood event likely eliminated the
recruitment of most, if not all, of those seedlings into the
population. At
[[Page 47121]]
least 100 plants were present at this site in 2012 (Homoya, pers.
comm., November 2012); however, it is not known whether these were
survivors of the flood or new plants that had sprouted from the seed
bank.
There are seven Tennessee occurrences that TDEC (2009, p. 2)
reported could be affected by water level manipulation. One of these,
Tennessee EO 3, is located on a wooded slope above the upper reaches of
waters impounded by Old Hickory Lake. There were more than 500 plants
at this location in 2008, and the position of Short's bladderpod within
the forested area above the zone of routine water level fluctuation is
unlikely to be affected by manipulation of water levels in the lake.
Shea (1993, p. 90) did not mention water level manipulation in her
assessment of threats to this occurrence. Tennessee EO 20, also in the
upper reaches of Old Hickory Lake, is presumed extirpated but was
likely lost to placement of rip rap along the roadside where it occurs,
as discussed above (please see Transportation Right-of-Way Construction
and Maintenance). Tennessee EO 12 is located on bluffs overlooking the
Cumberland River but not within an area managed as a reservoir or lake.
Shea (1993, pp. 22-23) was unable to find this occurrence in 1992, and
concluded that flooding at the base of the bluff was the cause. In
2008, TDEC (2009, p. 8) found approximately 50 plants at Tennessee EO
12, but they considered Short's bladderpod habitat to be vulnerable to
flooding at this site due to water level fluctuation and the position
of the plants at a low elevation on the bluff. Tennessee EOs 24 through
27 are found in soil at the river bank or on bedrock ledges within
about 1.5 m (5 ft) of the waters of Cordell Hull Reservoir (TNHID
2012), but, with the exception of EO 27, no more than 10 plants have
ever been counted at any of these sites. These three occurrences are
vulnerable to the effects of water level fluctuation, as evidenced by
observed erosion within the fluctuation zone (TNHID 2012). Tennessee EO
27 appears to be at little risk of habitat alteration due to water
level fluctuation, as it is located on bluff ledges above the zone of
routine water level fluctuation.
While the threat of flooding or water level fluctuation is present
at only five extant occurrences (19 percent), one of these is the only
Indiana population of the species, where the species has numbered in
excess of 1,000 plants in the past (Homoya, pers. comm., November
2012). The four occurrences in Tennessee threatened by water level
fluctuation are small and vulnerable to extirpation from even limited
habitat alteration or inundation.
Overstory Shading
The most vigorous (Shea 1992, p. 24) and stable (TDEC 2009, p. 1)
Short's bladderpod occurrences are found in locations where the canopy
has remained relatively open over time. Overstory shading appears to
have been a factor contributing to the disappearance of Short's
bladderpod at three sites in Kentucky (EO numbers 9, 19, and 20) and
one in Tennessee (EO 2) where Shea (1992, p. 4) observed heavy shading
as a threat to the species in 1992. Overstory shading has been
identified as a threat to Indiana EO 1 (INHDC 2012), Kentucky EO 22
(KNHP 2012), and Tennessee EOs 10, 21, and 24 (TNHID 2012), or 19
percent of known extant occurrences. Based on these data, canopy
shading has been implicated as a factor contributing to the
disappearance of Short's bladderpod from four sites and has been
identified as a limiting factor at nearly one-fifth of remaining extant
occurrences.
Competition With Nonnative Plant Species
Competition with or shading from invasive, nonnative herbaceous and
shrub species are cited in notes concerning threats in database records
for three of Kentucky's (EO numbers 4, 11, and 18) (KNHP 2012) and five
of Tennessee's (EO numbers 8, 10, 22, 24, and 26) (TNHID 2012) extant
Short's bladderpod occurrences. Homoya (pers. comm., December 2012)
also lists invasive species among the threats affecting the single
Indiana occurrence. The species most often mentioned by these agencies
include Lonicera japonica (Japanese honeysuckle), L. maackii (bush
honeysuckle), Alliaria petiolata (garlic mustard), and Bromus tectorum
(downy brome grass); however, several other invasive, nonnative species
occur in sites where Short's bladderpod exists, including Ligustrum
spp. (privet), Rosa multiflora (multiflora rose), and Glechoma
hederacea (ground ivy). Competition with or shading from these species
adversely affects Short's bladderpod. While this threat has been
specifically noted at approximately one-third of Short's bladderpod
occurrences, it likely is more widespread among occurrences of the
species and has not been reported in database records.
Trash Dumping
Shea (1993, p. 22) identified three Short's bladderpod sites at
which trash dumping posed a threat (Kentucky EOs 1 and 19, Tennessee EO
20). The species is no longer found at two of these sites: Kentucky EO
19, where canopy shading has been implicated in the species' absence,
and Tennessee EO 20, where most of the habitat for the species has been
covered by rip-rap. While Short's bladderpod is presumed to be extant
at Kentucky EO 1, there was only one plant found at this site in 2009
(KNHP 2012). The species was first collected at this site in 1957, and
despite several site visits between then and 2009, only two plants were
seen there in 1992 (KNHP 2012). TDEC (2009, p. 3) lists trash dumping
as a general threat to Short's bladderpod, but provides no specific
information to support this conclusion.
Livestock Grazing
Livestock grazing historically presented a threat to Short's
bladderpod, but we are not aware of any threats currently posed by this
land use. In addition to potentially causing direct harm to or loss of
individual plants, livestock grazing on the steeply sloped sites where
Short's bladderpod typically occurs could increase soil erosion,
potentially uprooting individual plants and causing loss of the soil
seed bank. Shea (1993, p. 22) identified three Kentucky sites (EOs 9,
20, and 21) at which livestock (goats or cows) grazing posed a threat
to Short's bladderpod. None of these sites support the species today,
likely due to multiple factors that degraded the habitat at those
locations. In Tennessee, Shea (1993, p. 22) reported that EO numbers 15
and 21 were threatened by grazing. However, more recent data from TDEC
(TNHID 2012) indicate that Short's bladderpod has remained relatively
stable at these sites and grazing is not listed among threats observed
at these locations.
Commercial and Residential Construction
While TDEC (2009, p. 3) lists commercial and residential
construction among potential threats to Short's bladderpod, there is
little documentation of these impacts. Tennessee EO 31, which is based
on a single herbarium collection from 1979, was apparently lost due to
construction activities at its location within the city of Clarksville
(TNHID 2012). The only other reference we have found for this
particular threat was an observation by TDEC (TNHID 2012) that an area
in the vicinity of Tennessee EO 21 had been subdivided for residential
construction on the bluffs overlooking Old Hickory Lake. Construction-
related threats to Short's bladderpod could include direct destruction
of habitat and the plants found there or the indirect effects of
habitat alteration from sediment runoff
[[Page 47122]]
and encroachment of invasive, nonnative plant species from areas
disturbed during construction.
Climate Change
Our analyses under the Act include consideration of ongoing and
projected changes in climate. The terms ``climate'' and ``climate
change'' are defined by the Intergovernmental Panel on Climate Change
(IPCC). ``Climate'' refers to the mean and variability of different
types of weather conditions over time, with 30 years being a typical
period for such measurements, although shorter or longer periods also
may be used (IPCC 2007a, p. 78). The term ``climate change'' thus
refers to a change in the mean or variability of one or more measures
of climate (e.g., temperature or precipitation) that persists for an
extended period, typically decades or longer, whether the change is due
to natural variability, human activity, or both (IPCC 2007, p. 78).
Various types of changes in climate can have direct or indirect effects
on species. These effects may be positive, neutral, or negative and
they may change over time, depending on the species and other relevant
considerations, such as the effects of interactions of climate with
other variables (e.g., habitat fragmentation) (IPCC 2007, pp. 8-14, 18-
19). In our analyses, we use our expert judgment to weigh relevant
information, including uncertainty, in our consideration of various
aspects of climate change.
The Intergovernmental Panel on Climate Change (IPCC) concluded that
evidence of warming of the climate system is unequivocal (IPCC 2007a,
p. 30). Numerous long-term climate changes have been observed including
changes in arctic temperatures and ice, widespread changes in
precipitation amounts, ocean salinity, wind patterns and aspects of
extreme weather including droughts, heavy precipitation, heat waves and
the intensity of tropical cyclones (IPCC 2007b, p. 7). While continued
change is certain, the magnitude and rate of change is unknown in many
cases. Species that are dependent on specialized habitat types, are
limited in distribution, or have become restricted to the extreme
periphery of their range will be most susceptible to the impacts of
climate change.
Estimates of the effects of climate change using available climate
models lack the geographic precision needed to predict the magnitude of
effects at a scale small enough to discretely apply to the range of
Short's bladderpod. However, data on recent trends and predicted
changes for the Southeast and Midwest United States (Karl et al. 2009,
pp. 111-122) provide some insight for evaluating the potential threat
of climate change to the species. Most of the range of Short's
bladderpod lies within the geographic area included by Karl et al.
(2009, pp. 111-122) in their summary of regional climate impacts
affecting the Southeast region; however, the Indiana occurrence of the
species lies in the Midwest region, just west of its boundary with the
Southeast region.
Since 1970, the average annual temperature across the Southeast has
increased by about 2 [deg]F, with the greatest increases occurring
during winter months. The geographic extent of areas in the Southeast
region affected by moderate to severe spring and summer drought has
increased over the past three decades by 12 and 14 percent,
respectively (Karl et al. 2009, p. 111). These trends are expected to
increase. Rates of warming are predicted to more than double in
comparison to what the Southeast has experienced since 1975, with the
greatest increases projected for summer months. Depending on the
emissions scenario used for modeling change, average temperatures are
expected to increase by 4.5 [deg]F to 9 [deg]F by the 2080s (Karl et
al. 2009, p. 111). While there is considerable variability in rainfall
predictions throughout the region, increases in evaporation of moisture
from soils and loss of water by plants in response to warmer
temperatures are expected to contribute to increased frequency,
intensity, and duration of drought events (Karl et al. 2009, p. 112).
Projected increases in winter and spring rainfall for the Midwest
region, as well as predictions of more intense rainfall events
throughout the year, are expected to lead to more frequent flooding.
Despite these projected trends, the likelihood of drought is expected
to increase in the Midwest due to warming-induced increases in
evapotranspiration rates and longer intervals between precipitation
events (Karl et al. 2009, pp. 120-121).
Depending on timing and intensity of drought events, Short's
bladderpod could be adversely affected by increased mortality rates,
reduced reproductive output due to loss or reduced vigor of mature
plants, and reduced rates of seed germination and seedling recruitment.
The species' presumed ability to form a seed bank should provide some
resilience to drought-induced population declines; however, multiple
droughts in successive years could diminish this resilience and lead to
the loss of occurrences. Conversely, increased drought frequency and
severity could alter structure of vegetation communities in which
Short's bladderpod occurs by slowing rates of forest canopy
development, increasing tree mortality, and increasing light
availability for the species, which could stimulate recruitment from
dormant seed banks and increase vigor of plants located in areas that
are presently well-shaded. The predicted increase in flood frequency in
the Midwest could place the Indiana population of the species at risk,
as evidenced by the loss of large numbers of seedlings during a
prolonged flood at this site in 2011. While climate has changed in
recent decades in regions where Short's bladderpod occurs and the rate
of change likely will continue to increase into the future, we do not
have data to determine how the habitats where Short's bladderpod occurs
will be affected by these changes and how the species will respond to
these changes.
Conservation Efforts To Reduce Habitat Destruction, Modification, or
Curtailment of Its Range
There have been limited conservation efforts directed towards
reducing threats affecting Short's bladderpod and its habitat. The
Indiana Department of Natural Resources acquired the single Indiana
occurrence. IDNR controls competing vegetation by mowing along the
roadside where Short's bladderpod occurs and attempts to stimulate
germination and seedling recruitment with light soil disturbance. The
species has responded positively, at least in the short term, to this
management (Homoya, pers. comm., December 2012). In Kentucky, a
Landowner Incentive Program grant was used to manage vegetation
structure or control invasive species at two occurrences in 2005. The
effort to control bush honeysuckle at Kentucky EO 19 provided only a
short-term benefit, if any, for Short's bladderpod, as bush honeysuckle
is again well established at this site. During 2011, no Short's
bladderpod plants could be found at this site, and the occurrence is
presumed extirpated. The removal of cedar trees at Kentucky EO 23
appears to have positively affected habitat conditions for Short's
bladderpod, as there were more than 500 plants, mostly seedlings,
observed at the site in 2011. The Kentucky State Nature Preserve
Commission acquired lands to establish the Rockcress Hills State Nature
Preserve, where Kentucky EO 22 is located and where the federally
listed endangered Braun's rockcress (listed as Arabis perstellata, but
now recognized as Boechera perstellata) also occurs. As discussed
above, this occurrence is threatened by shading due to forest canopy
development. These conservation efforts have benefited three
[[Page 47123]]
extant Short's bladderpod occurrences, but significant habitat threats
remain across the species' range.
Summary of Factor A
The threats to Short's bladderpod from habitat destruction and
modification are occurring throughout the entire range of the species.
These threats include transportation right-of-way construction and
maintenance; flooding and water level fluctuation; overstory shading;
and competition with nonnative plant species. The population level
impacts from these activities are expected to continue into the future.
Trash dumping, livestock grazing, and commercial and residential
construction have been recognized as threats to habitat for this
species, but there is little evidence that these are significant
threats to extant occurrences.
B. Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
There has been limited collection of Short's bladderpod seed for
conservation purposes. The Missouri Botanical Garden holds seed
accessions from the Indiana occurrence, four Kentucky occurrences (EOs
4, 18, 19, and 28), and two Tennessee occurrences (EOs 4 and 17).
Kentucky EO 19 is no longer extant, for reasons discussed above, but
Short's bladderpod is still found at all of the other occurrences from
which these accessions were collected. Dr. Carol Baskin (pers. comm.,
December 2012) collected seeds from Indiana for research on seed
ecology. We are not aware of commercial trade in Short's bladderpod at
this time. Indiscriminate collecting for scientific or other purposes
could be a threat to the species due to the low numbers of individuals
at most occurrences, but we have no data to indicate that
indiscriminate collecting of Short's bladderpod has occurred. On the
contrary, collections for ex situ conservation holdings could be an
important component of future recovery efforts for the species.
C. Disease or Predation
We are not aware of any commercial or scientific data indicating
that disease or predation threatens the continued existence of Short's
bladderpod.
D. The Inadequacy of Existing Regulatory Mechanisms
Section 4(b)(1)(A) of the Act requires the Service to take into
account ``those efforts, if any, being made by any State or foreign
nation, or any political subdivision of a State or foreign nation, to
protect such species. . . .'' In relation to Factor D under the Act, we
interpret this language to require the Service to consider relevant
Federal, State, and tribal laws, plans, regulations, and other such
mechanisms that may minimize any of the threats we describe in threat
analyses under the other four factors, or otherwise enhance
conservation of the species. We give strongest weight to statutes and
their implementing regulations and to management direction that stems
from those laws and regulations. An example would be State governmental
actions enforced under a State statute or constitution, or Federal
action under statute.
Having evaluated the significance of the threat as mitigated by any
such conservation efforts, we analyze under Factor D the extent to
which existing regulatory mechanisms are inadequate to address the
specific threats to the species. Regulatory mechanisms, if they exist,
may reduce or eliminate the impacts from one or more identified
threats. In this section, we review existing State and Federal
regulatory mechanisms to determine whether they effectively reduce or
remove threats to Short's bladderpod.
Short's bladderpod is listed as endangered in Indiana, Kentucky,
and Tennessee. In Indiana this listing does not provide legal
protection for the species; although, listed species are given special
consideration when planning government-funded projects. Additionally,
the Indiana site is located on land owned by the IDNR where collection
or damage to plants is prohibited.
The Kentucky Rare Plants Recognition Act, Kentucky Revised Statutes
(KRS), chapter 146, section 600-619, directs the KSNPC to identify
plants native to Kentucky that are in danger of extirpation within
Kentucky and report every 4 years to the Governor and General Assembly
on the conditions and needs of these endangered or threatened plants.
This list of endangered or threatened plants in Kentucky is found in
the Kentucky Administrative Regulations, title 400, chapter 3:040. The
statute (KRS 146:600-619) recognizes the need to develop and maintain
information regarding distribution, population, habitat needs, limiting
factors, other biological data, and requirements for the survival of
plants native to Kentucky. This statute does not include any regulatory
prohibitions of activities or direct protections for any species
included in the list. It is expressly stated in KRS 146.615 that this
list of endangered or threatened plants shall not obstruct or hinder
any development or use of public or private land. Furthermore, the
intent of this statute is not to ameliorate the threats identified for
the species, but it does provide information on the species.
The Tennessee Rare Plant Protection and Conservation Act of 1985
(T.C.A. 11-26-201) authorizes the Tennessee Department of Environment
and Conservation (TDEC) to, among other things: conduct investigations
on species of rare plants throughout the state of Tennessee; maintain a
listing of species of plants determined to be endangered, threatened,
or of special concern within the state; and regulate the sale or export
of endangered species via a licensing system. This act forbids persons
from knowingly uprooting, digging, taking, removing, damaging,
destroying, possessing, or otherwise disturbing for any purpose, any
endangered species from private or public lands without the written
permission of the landowner, lessee, or other person entitled to
possession and prescribes penalties for violations. The TDEC may use
the list of threatened and special concern species when commenting on
proposed public works projects in Tennessee, and the department shall
encourage voluntary efforts to prevent the plants on this list from
becoming endangered species. This authority shall not, however, be used
to interfere with, delay, or impede any public works project.
Thus, despite the fact that Short's bladderpod is listed as
endangered by the states of Indiana, Kentucky, and Tennessee, these
designations confer no guarantee of protection to the species or its
habitat, whether on privately owned or state-owned lands, unless such
protections are voluntarily extended to the species.
E. Other Natural or Manmade Factors Affecting Its Continued Existence
The ability of populations to adapt to environmental change is
dependent upon genetic variation, a property of populations that
derives from its members possessing different forms (i.e., alleles) of
the same gene (Primack 1998, p. 283). Small populations occurring in
isolation on the landscape can lose genetic variation due to the
potentially strong influence of genetic drift, i.e., the random change
in allele frequency from generation to generation (Barrett and Kohn
1991, p. 8). Smaller populations experience greater changes in allele
frequency due to drift than do larger populations (Allendorf and
Luikart 2007, pp. 121-122). Loss of genetic variation due to genetic
drift heightens susceptibility of small populations to adverse genetic
effects, including inbreeding depression and
[[Page 47124]]
loss of evolutionary flexibility (Primack 1998, p. 283). Deleterious
effects of loss of genetic variation through drift have been termed
drift load, which is expressed as a decline in mean population
performance of offspring in small populations (Willi et al. 2005, p.
2260).
The likelihood that Short's bladderpod is self-incompatible
presents another threat related to small population sizes. Genetic
incompatibility prevents self-fertilization or reduces successful
breeding among closely related individuals, which can decrease mean
fitness in small populations because of increased probability of an
encounter of two incompatible haplotypes (specific combination of
alleles at adjacent locations (loci) on the chromosome that are
inherited as a unit) (Willi et al. 2005, p. 2256), which would prevent
seed production in self-incompatible plants. In small populations, less
common S-haplotypes (self-incompatibility haplotypes) might be easily
lost due to genetic drift, reducing the number of compatible mates
within the population (Byers and Meagher 1992, p. 356).
In self-incompatible plants of the Brassicaceae family, when pollen
and stigma share S-haplotypes at the S-locus (self-incompatibility
locus, i.e., the position on a chromosome occupied by the self-
incompatibility gene complex), pollen tube development is disrupted on
the stigma of the female reproductive system (Takayama and Isogai 2005,
p. 469). The stigma is the receptive structure of the female
reproductive system in plants, which also includes the pistil and
ovary, on which pollen grains germinate and begin development of the
pollen tube. Pollen tube formation is necessary for fertilization of
the ovary and subsequent seed production to occur.
Despite the presence of such a mechanism functioning to reduce or
eliminate reproductive output among individuals sharing S-haplotypes,
in small populations mating is likely to occur among individuals that
possess different S-haplotypes but are genetically similar at other
loci due to loss of alleles from the population through genetic drift
(Byers and Meagher 1992, p. 358). Mating between such closely related
individuals is referred to as inbreeding. Inbreeding rates are higher
in small populations because most or all individuals in the population
are related, and inbred individuals generally have reduced fitness as
compared to non-inbred individuals from the same population, a
phenomenon referred to as inbreeding depression (Allendorf and Luikart
2007, p. 306).
Evidence in plants of inbreeding depression due to small population
size is provided by Heschel and Paige (1995, p. 128), who found that
plants from populations of Ipomopsis aggregata (scarlet gilia) with 100
or fewer flowering individuals produced smaller seeds with lower rates
of germination success compared to those from populations with more
than 100 flowering individuals. Heschel and Paige (1995, p. 131) also
found that seed sizes increased and germination success improved in
response to transfer of pollen into each of the small populations,
which they interpreted as evidence that the reduced fitness observed in
small populations was attributable, in part, to inbreeding depression.
Willi et al. (2005, pp. 2263) found evidence of the three processes
described above (reduced cross-compatibility presumably due to lack of
compatible mates carrying different S-haplotypes, reduced fitness due
to inbreeding, and drift load due to loss of genetic variation)
simultaneously affecting small populations of a plant, Ranunculus
reptans (creeping buttercup). Populations with low allelic diversity,
taken as an indication of long-term small population size, had higher
inbreeding levels. Inbreeding depression in these populations was
expressed as poor clonal performance and reduced seed production in
offspring (F1 plants) produced by crosses between plants with high
kinship coefficients. Drift load also was expressed as a reduction in
mean seed production of F1 plants in long-term small populations (Willi
et al. 2005, p. 2260).
In evaluating threats to Short's bladderpod that could arise due to
small population size, we first evaluated the limited data available
concerning abundance at each of the occurrences across the species'
range. This represents a conservative classification of small
population size, as available data typically do not discriminate among
life history stages, so the number of reproducing individuals is
typically less than what is shown in the abundance data in Table 1 (see
Distribution and Status for the Short's bladderpod, above). Less than
100 individual plants have ever been observed at one time at 12 (46
percent) of the extant occurrences in Kentucky (EOs 1, 11, and 28) and
Tennessee (EOs 8, 12, 15, 22, 24, 26, 27, 29, and 30). The greatest
number of plants ever observed at the small Kentucky occurrences ranged
from 2 at EO 1 to 52 at EO 11 (KNHP 2012). At the small Tennessee
occurrences, maximum recorded abundance ranged from 3 clusters of
plants at EO 26 to approximately 50 plants each at EOs 8, 12, 22, 27,
and 29 (TNHID 2012). These small populations are at risk of adverse
effects from reduced genetic variation and associated drift load,
increased risk of inbreeding depression, and reduced reproductive
output due to low availability of genetically compatible mates. Many of
these occurrences where population sizes are small are isolated from
other occurrences, decreasing the likelihood that they could be
naturally reestablished via seed dispersal, in the event that local
extinction occurred.
Cumulative Effects From Factors A through E
Where two or more threats affect Short's bladderpod occurrences,
the effects of those threats could interact or be compounded, producing
a cumulative adverse effect that rises above the incremental effect of
either threat alone. The most obvious cases in which cumulative adverse
effects would be significant are those in which small populations
(Factor E) are affected by threats that result in destruction or
modification of habitat (Factor A). Two occurrences in Kentucky and six
in Tennessee where small population size was identified as a threat
also face threats to their habitats, as discussed under Factor A above.
The vulnerability of these occurrences to habitat modification or
destruction is heightened by effects of small population size discussed
above, reduced resilience to recover from acute demographic effects of
habitat disturbances, and low potential for recolonization due to
isolation from other occurrences.
Whorled Sunflower
A. The Present or Threatened Destruction, Modification, or Curtailment
of Its Habitat or Range
Whorled sunflower appears to be a narrow habitat specialist,
occurring in natural wet meadows or prairies and calcareous barrens.
Such habitats likely were more extensive in the eastern United States
before European settlement, subsequent fire suppression, and conversion
of habitat to cropland or residential areas (Allison 1995, p. 7). Today
these prairie areas are not very extensive, and they often are degraded
or have been destroyed for a number of reasons. Most remaining prairie
vegetation in the geographic area where whorled sunflower occurs exists
as remnants along roadside and utility rights-of-way, where prairie-
like
[[Page 47125]]
conditions are artificially maintained (Allison 1995, p. 4). Where
whorled sunflower habitat remains, it faces threats due to
indiscriminate use of mechanical or chemical vegetation management for
industrial forestry, right-of-way maintenance, or agricultural purposes
that could adversely affect it. Because the species requires well-lit
habitats for its growth and reproduction, shading and competition due
to vegetation succession in the absence of natural or human-caused
disturbance also threaten whorled sunflower habitat.
Industrial Forestry Practices
Industrial forestry practices have altered much suitable whorled
sunflower habitat in Georgia and Alabama, and currently threaten one
known subpopulation in Alabama. While surveying potential habitat for
additional populations, J. Allison (Botanist, Georgia Department of
Natural Resources, pers. comm., March 1999) observed that much of this
species' prairie habitat in Georgia had been converted to pine
plantations. Nearly all of the Georgia subpopulations and one of the
Alabama subpopulations of whorled sunflower are located on lands that
currently are owned by The Campbell Group, a timberland investment
advisory firm. The Georgia subpopulations on The Campbell Group's lands
are protected from habitat destruction or degradation by their
inclusion in the conservation easement area at the Coosa Valley
Prairie, which was donated to The Nature Conservancy by the Temple-
Inland Corporation, the former owner of these lands.
With the exception of the conservation easement area at the Coosa
Valley Prairie, The Campbell Group typically subsoil plows planting
sites to improve drainage and conditions for tree root development, and
uses mechanical or chemical methods to control competing vegetation
when preparing sites for planting pine seedlings (J. King, Area
Manager, The Campbell Group, LLC, pers. comm., August 2012) on its
lands in Floyd County, Georgia, and Cherokee County, Alabama. These
practices could cause direct mortality of whorled sunflower plants at
one of the Alabama subpopulations and could contribute to habitat
degradation caused by shading and competition (please see ``Shading and
Competition'' below) by improving conditions for growth of planted
pines. During timber harvests, either to thin (i.e., reduce density of
pine trees in order to improve growth conditions for remaining trees)
or to clearcut the stand, whorled sunflower plants at this
subpopulation could be subjected to indirect adverse effects from soil
disturbance or to direct mortality due to movement of harvesting
equipment.
Right-of-Way Maintenance
Incompatible maintenance activities in transportation rights-of-way
have adversely affected the whorled sunflower in Alabama and Tennessee,
and could affect one subpopulation in Georgia. At one of the Alabama
subpopulations, the whorled sunflower occurs in a narrow strip of
vegetation between a roadside and adjacent pine forest, where it is
vulnerable to mortality or reduced vigor and reproductive output due to
indiscriminate use of herbicides or mowing for right-of-way
maintenance. Poorly timed mowing of this right-of-way prevented
flowering and seed production in some plants at this site in 2008;
however, the Alabama Department of Conservation and Natural Resources,
Alabama Department of Transportation, and Cherokee County Highway
Department cooperated in placing signs at the site to mark the presence
of whorled sunflower and to attempt to prevent this in the future (W.
Barger, Botanist, Alabama Department of Conservation and Natural
Resources, pers. comm., February 2009); periodic replacement might be
needed due to vandalism or removal of the signs (Barger, pers. comm.,
March 2012). Regular coordination with parties responsible for roadside
maintenance at this location will be necessary to avoid future adverse
effects to the whorled sunflower from indiscriminate mowing or
herbicide application.
Plants extending onto a roadside within a powerline right-of-way at
the Madison County, Tennessee, population were subjected to herbicide
spraying in association with roadside and powerline maintenance in
2004, causing significant mortality (A. Bishop, Botanist, TDEC, pers.
comm., February 2008; D. Lincicome, Natural Heritage Program Manager,
TDEC, pers. comm., September 2006). Similarly, plants extending into
the railroad right-of-way at the McNairy County, Tennessee, population
are vulnerable to adverse effects from indiscriminate herbicide
application for railroad right-of-way maintenance. A small cluster of
plants in one of the Georgia's subpopulations is located on the bank of
a road adjacent to the Coosa Valley Prairie easement area and is not
protected. These data indicate that effects of indiscriminate use of
herbicides or mowing for vegetation management in transportation
rights-of-way could adversely affect the whorled sunflower populations
in Alabama and Tennessee, as well as a small subpopulation in Georgia.
Agricultural Practices and Land Conversion
The whorled sunflower has not been rediscovered at the type
locality in Tennessee despite intensive surveys of that area (Nordman
1998, p. 1-2). However, this record is from an 1892 collection and
locality information is vague, so it is not possible to determine why
this population has been lost. In Tennessee, much of this species'
suitable habitat presumably has been converted for agricultural use, as
substantial proportions of the counties in the State where the species
have been found have been in row crop production since 1850 (Table 5)
(Waisanen and Bliss 2002; GIS data available at https://landcover.usgs.gov/cropland, accessed January 9, 2013). Because this
species was not seen following the initial 1892 collection until it was
rediscovered in 1994, and was not seen again in Tennessee until 1998,
it is impossible to know the historical distribution and abundance of
its habitat. However, the data in Table 5 indicate that land conversion
to agricultural uses has a long and sustained history in the Tennessee
counties where the whorled sunflower has been found and likely has
contributed to loss of habitat and whorled sunflower populations.
Table 5--Proportions of County Land Base Considered Improved Farmland for Tennessee Counties Where the Whorled
Sunflower Has Been Found. Reported Here for Each County Are the Highest and Lowest Proportions on Record for
Each County and the Years in Which They Occurred and Values for the Years 1850 and 1997, the First and Last
Years Included in Waisanen and Bliss (2002).
----------------------------------------------------------------------------------------------------------------
County High (year) Low (year) 1850 1997
----------------------------------------------------------------------------------------------------------------
Chester..................................................... 37 (1940) 18 (1850) 18 23
Madison..................................................... 54 (1949) 23 (1870) 28 29
[[Page 47126]]
McNairy..................................................... 33 (1920) 14 (1850) 14 20
----------------------------------------------------------------------------------------------------------------
Agricultural practices, including field preparation, herbicide use,
and harvesting of crops, are threats to both of the known Tennessee
populations, due to the species' presence in habitats adjacent to
actively farmed crop fields in both locations. In July 2009, TDEC
biologists observed that one clump consisting of two whorled sunflower
stems had been destroyed by row crop cultivation in a previously fallow
field at the McNairy County, Tennessee, population. Unpaved access
roads around the perimeter of this field had also been widened,
encroaching on whorled sunflower plants (7 clumps, 140 stems) in an
adjacent railroad right-of-way (Bishop, pers. comm., March 2010). With
the exception of the approximately 1-ha (2.5-ac) patch of old field
habitat discussed above (see Habitat for the whorled sunflower, above),
the Madison County, Tennessee, whorled sunflower population is
distributed in narrow strips of vegetation along borders of row crop
fields and is vulnerable to mechanized disturbance of these habitats or
to effects from herbicide application. Based on this information we
conclude that habitat at both whorled sunflower populations in
Tennessee face significant threats associated with agricultural
practices used in row crop production.
Shading and Competition
Absent natural or human-caused disturbance, habitats where whorled
sunflower occurs are threatened by succession of vegetation to a shrub-
dominated or forested condition. The largest concentration of plants at
the Madison County, Tennessee, population is located in a successional
old field approximately 1 ha (2.5 ac) in size, where vegetation
succession threatens to degrade the largest patch of contiguous habitat
where the majority of this population occurs. Woody species present at
this site include Acer negundo (box elder), Liquidambar styraciflua
(sweetgum), and Salix nigra (black willow) (Tennessee Division of
Natural Areas 2006, p. 5), all of which can rapidly invade moist old
field habitats if left unmanaged. No conservation agreements or
management plans are in place to ensure that this site receives
periodic disturbance to maintain open conditions needed for the growth
and sexual reproduction of whorled sunflower.
The Alabama subpopulation on The Campbell Group's lands is located
in a site where the prior owner, Temple-Inland Corporation, harvested
an immature hardwood forest in 1998. Initially this timber harvest was
thought to have adversely affected the whorled sunflower population,
but these plants and associated prairie species responded favorably
within a few years following the harvest. However, the site was
subsequently converted into a loblolly pine plantation, and the trees
have attained sufficient size and density to threaten whorled sunflower
plants due to increased shading and competition (Schotz 2011, p. 4). As
of 2012, there were few whorled sunflower plants present at this site,
and those present were in a suppressed, vegetative condition due to
strong shading and competition from planted pines and vegetation
growing in the understory. Encroachment by invasive, nonnative plants
following the timber harvest and establishment of the loblolly pine
stand also is a threat at this site (Schotz 2011, p. 12). The second
Alabama subpopulation is relegated to a narrow strip of vegetation
between a roadside and adjacent pine forest with a densely vegetated
understory. The spatial extent of this subpopulation is limited by the
whorled sunflower's inability to grow in the shaded habitat of the
adjacent forest.
Based on this information we conclude that habitat degradation due
to shading and competition resulting from vegetation succession
currently is a significant threat to two whorled sunflower populations.
Both of the Alabama subpopulations and the largest contiguous patch of
suitable occupied habitat for the species in Tennessee are at risk from
this threat.
Climate Change
We discuss the topic of climate change in greater detail above in
the Factor A threats analysis for Short's bladderpod, which is also
applicable to whorled sunflower. Since 1970, the average annual
temperature across the Southeast has increased by about 2 [deg]F, with
the greatest increases occurring during winter months. The geographic
extent of areas in the Southeast region affected by moderate to severe
spring and summer drought has increased over the past three decades by
12 and 14 percent, respectively (Karl et al. 2009, p. 111). These
trends are expected to increase. Rates of warming are predicted to more
than double in comparison to what the Southeast has experienced since
1975, with the greatest increases projected for summer months.
Depending on the emissions scenario used for modeling change, average
temperatures are expected to increase by 4.5 [deg]F to 9 [deg]F by the
2080s (Karl et al. 2009, p. 111). While there is considerable
variability in rainfall predictions throughout the region, increases in
evaporation of moisture from soils and loss of water by plants in
response to warmer temperatures are expected to contribute to increased
frequency, intensity, and duration of drought events (Karl et al. 2009,
p. 112).
The predicted increase in drought frequency, intensity, and
duration could adversely affect the moist prairie habitats inhabited by
whorled sunflower, by reducing soil moisture and increasing sunflower
mortality rates or reducing flowering and seed production rates. A
positive effect of increased drought could result from increased
mortality of woody vegetation and reduced rates of vegetation
succession, which diminishes habitat abundance and quality for whorled
sunflower. While climate has changed in recent decades in the region
where whorled sunflower occurs and the rate of change likely will
continue to increase into the future, we do not have data to determine
how the habitats where the whorled sunflower occurs will be affected by
these changes and how the species will respond to these changes.
Conservation Efforts To Reduce Habitat Destruction, Modification, or
Curtailment of Its Range
Temple-Inland Corporation donated a conservation easement for the
Coosa Valley Prairie property in Georgia to The Nature Conservancy,
thereby
[[Page 47127]]
protecting most of the Georgia population of this species. This site
drains into the headwaters of Mud Creek. In 2002, The Georgia
Department of Natural Resources and The Nature Conservancy worked with
staff of Temple-Inland to develop a 10-year management plan for
conservation of rare species within this easement area. Site-specific
management plans for several open wet prairies, known to provide
habitat for this species within the easement, were developed. Temple-
Inland implemented a prescribed burn and selective timber harvest on
243 ha (600 ac) of the easement in 2001, to improve habitat conditions
for whorled sunflower and other species. Temple-Inland conducted
additional burns within the easement area between 2002 and 2006.
Mechanical thinning and control of invasive, exotic plants was also a
component of their management of this site.
This easement area, now owned by The Campbell Group, is
cooperatively managed with The Nature Conservancy based on a jointly
developed conservation management plan, which was revised in 2012, for
the period extending through 2016. The management goals for the site
are based on the conservation easement and include long-term
perpetuation and restoration of the mosaic of prairies, woodlands,
wetlands, creeks, and forest while allowing for sustainable timber
harvesting. Protecting and enhancing native plant communities,
especially those supporting rare species, is the primary management
objective, and periodic timber harvesting is a secondary objective.
Portions of the tract either have been or will be planted into Pinus
palustris (longleaf pine) as part of the Longleaf Alliance partnership.
Prescribed fire is the primary management tool used to perpetuate and
restore the native plant communities and also serves silvicultural
objectives.
Despite the existence of a conservation plan and the cooperative
partnership between The Nature Conservancy and The Campbell Group to
implement the plan, management with prescribed fire is not a binding
condition of the conservation easement. Thus, the potential remains
that this management could be discontinued in the event that the
property was sold to a less cooperative landowner.
Summary of Factor A
The threats to whorled sunflower from habitat destruction and
modification are occurring throughout the entire range of the species.
These threats include mechanical or chemical vegetation management
associated with industrial forestry practices, maintenance of
transportation and utility rights-of-way, agricultural practices, and
shading and competition. While a conservation easement and suitable
habitat management alleviate threats from industrial forestry that
otherwise would adversely affect the Georgia population, one of the
Alabama whorled sunflower subpopulations currently is threatened by
industrial forestry practices. The population-level impacts from these
activities are expected to continue into the future.
B. Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
The whorled sunflower currently is of limited availability in the
horticultural trade, although no negative impacts are known to have
occurred due to collection of wild material for commercial sale.
Nonetheless, the conspicuous, attractive flowers of this species
combined with easy access of some sites leaves the species vulnerable
to collection or poaching. Poaching from the small populations of
whorled sunflower that are known to exist could contribute to altered
demographic or genetic structure of populations, potentially
diminishing their viability; however, we have no information to suggest
this currently is an active threat or has adversely affected
populations in the past.
C. Disease or Predation
We are not aware of any commercial or scientific data indicating
that disease or predation threatens the continued existence of whorled
sunflower.
D. The Inadequacy of Existing Regulatory Mechanisms
Whorled sunflower is State-listed as endangered in Georgia and
Tennessee, but has no official State status in Alabama. The law that
provides official protection to designated species of plants in Georgia
is known as the Wildflower Preservation Act of 1973 (O.C.G.A. 12-6-
170). Under this law, no protected plant may be collected without
written landowner permission. No protected plant may be transported
within Georgia without a transport tag with a permit number affixed.
Permits are also used to regulate a wide array of conservation
activities, including plant rescues, sale of protected species, and
propagation efforts for augmentation of natural populations and
establishment of new ones. No protected plants may be collected from
State-owned lands without the express permission of the Georgia
Department of Natural Resources. The Georgia Environmental Policy Act
(GEPA; O.C.G.A. 12-16-1), enacted in 1991, requires that impacts to
protected species be addressed for all projects on State-owned lands,
and for all projects undertaken by a municipality or county if funded
half or more by State funds, or by a State grant of more than $250,000.
The provisions of GEPA do not apply to actions of nongovernmental
entities. On private lands, the landowner has ultimate authority over
what protection efforts, if any, occur with regard to protected plants
(Patrick et al. 1995, p. 1 of section titled ``Legal Overview'').
The Tennessee Rare Plant Protection and Conservation Act of 1985
(T.C.A. 11-26-201) authorizes the Tennessee Department of Environment
and Conservation (TDEC) to, among other things: conduct investigations
on species of rare plants throughout the state of Tennessee; maintain a
listing of species of plants determined to be endangered, threatened,
or of special concern within the state; and regulate the sale or export
of endangered species via a licensing system. This act forbids persons
from knowingly uprooting, digging, taking, removing, damaging,
destroying, possessing, or otherwise disturbing for any purpose, any
endangered species from private or public lands without the written
permission of the landowner, lessee, or other person entitled to
possession and prescribes penalties for violations. The TDEC may use
the list of threatened and special concern species when commenting on
proposed public works projects in Tennessee, and the department shall
encourage voluntary efforts to prevent the plants on this list from
becoming endangered species. This authority shall not, however, be used
to interfere with, delay, or impede any public works project.
Thus, despite the fact that whorled sunflower is listed as
endangered by the states of Georgia and Tennessee, these designations
confer no guarantee of protection to the species or its habitat,
whether on privately owned or state-owned lands, unless such
protections are voluntarily extended to the species by owners or
managers of lands where the species is present.
E. Other Natural or Manmade Factors Affecting Its Continued Existence
The whorled sunflower is vulnerable to localized extinction because
of its extremely restricted distribution and small population sizes at
most known locations, which reduces the resilience of these populations
to recover from acute demographic effects of threats to its habitat
discussed above under Factor A. Whorled sunflower is dependent
[[Page 47128]]
upon existence of prairie-like openings or remnant roadside prairie
habitats for its survival. Alteration or elimination of disturbance
processes that maintain these openings could result in the extinction
of populations of this species. Further, the highly fragmented
distribution of populations within Tennessee, combined with their
disjunct location with respect to those in Georgia and Alabama,
presumably precludes gene flow among them and leaves little chance of
natural recolonization of these populations in the event of localized
extinctions.
Small population size could be affecting reproductive fitness of
the whorled sunflower. The findings of Ellis and McCauley (2008,
entire) suggest that the Madison County, Tennessee, population is
reproductively less fit than the Alabama population. Ellis and McCauley
(2008, p. 1840) offered two possible explanations for reduced
reproductive fitness of the Tennessee population, including limited
mate availability due to limited diversity of self-incompatibility
alleles, or more extensive inbreeding. Both could be contributing to
reduced seed production and viability rates.
Ellis and McCauley (2008, pp. 1837-1838) could not assess the
fitness of the Georgia population because seed heads collected for the
study contained very few viable achenes, which produced poor
germination rates. However, the lack of viable achenes in seed heads
collected for this study suggests that poor reproductive fitness could
be a threat in this population, as well.
Cumulative Effects From Factors A through E
Where two or more threats affect whorled sunflower populations, the
effects of those threats could interact or be compounded, producing a
cumulative adverse effect that rises above the incremental effect of
either threat alone. Cumulative adverse effects are likely significant
for whorled sunflower because all of the populations are small and
their reproductive fitness is likely diminished (Factor E), and the
Alabama and Tennessee populations are affected by threats that result
in destruction or modification of habitat (Factor A). The vulnerability
of these occurrences to habitat modification or destruction is
heightened by the effects of small population size discussed above,
reduced resilience to recover from acute demographic effects of these
disturbances, and low potential for recolonization due to isolation
from other occurrences.
Fleshy-Fruit Gladecress
Factor A. The Present or Threatened Destruction, Modification, or
Curtailment of Its Habitat or Range
This species is endemic to cedar glade areas in north-central
Alabama that have been significantly altered from their original
condition. More than a 50 percent loss in glade habitat has occurred
since European settlement (Hilton 1997), with resulting glade habitats
reduced to remnants fragmented by agriculture and development. Hilton
(1997) conducted a thorough survey of cedar glade communities in
northern Alabama using historical records, soil maps, topographic maps,
geology, and aerial photography; 22 high priority glades were
identified. However, field surveys found only five of these to be in
good condition and restorable, and only two of these were considered
high-quality sites (Hilton, pers. comm., 1999).
Agricultural Practices
At four of the fleshy-fruit gladecress populations, plants occur in
pasture areas, on roadside rights-of-way, and/or in planted fields
surrounded by agriculture or residential developments (Hilton 1997, pp.
13-27). Periodic disturbance, such as plowing in row crop farming,
arrests succession and maintains populations in this type of habitat;
however, plowing or herbicide application in the spring prior to seed
set and dispersal could be detrimental to populations. Populations are
enhanced by disturbance created from light grazing, but heavy grazing
of pastures creates unfavorable conditions (i.e., soil compaction,
nutrient enrichment) for fleshy-fruit gladecress. Plants have been
severely trampled where grazing is allowed during the height of the
plant's flowering or fruiting period. Grazing during the reproductive
period also reduces vigor of the populations (Schotz, 2009, p. 2).
Improving pastures with fertilizer treatments or planting of forage
grasses could eventually result in loss of populations due to
competition. Lyons (in litt. 1981 to R. Sutter) considered that her
failure to relocate many of the historical fleshy-fruit gladecress
sites from the 1960s was due to the change in agricultural practices
from growing corn to using those sites for cattle pastures. McDaniel
and Lyons (1987, p. 11) considered the trend toward converting
agricultural sites from row crop cultivation to pasture as a primary
threat to the species.
Transportation Right-of-Way Maintenance
Five of the six fleshy-fruit gladecress occurrences extend onto
roadsides or are near roads, where mowing and herbicide application
prior to seed set pose threats to the species. Three historical sites
near roads have not been relocated and a portion of one of the extant
populations was destroyed by road widening and grading in the 1980s
(McDaniel and Lyons 1987, p. 7-9). Additional road widening at this
site in recent years has further reduced the size of this population
(Schotz 2009, p. 14). The largest population of this species has a dirt
road traversing through a portion of the site, which has made the site
vulnerable to off-road vehicles and dumping (Hilton 1997, p. 31). Other
sites have also been negatively affected by trash dumping and off-road
vehicles, including the site on U.S. Forest Service land. The U.S.
Forest Service has posted the area as closed and recently gated the
area to block all-terrain vehicle access to the site (T. Counts, U.S.
Forest Service, in litt. 2008), which appears to have been effective at
reducing damage to the glade (A. Cochran, U.S. Forest Service, in litt.
2005, Schotz in litt. 2007). The U.S. Forest Service continues to
monitor the glade site for impacts from recreational vehicles and from
other illegal vehicle activity (A. Cochran, pers. comm., 2011).
Shading and Competition
Winter annuals, such as fleshy-fruit gladecress, are excluded from
many habitats because they are poor competitors (Baskin and Baskin
1985, p. 387). As with all annuals, this species' long-term survival at
a locality is dependent upon its ability to reproduce and reseed there
every year. Thus, populations decline and become at risk of local
extinction if conditions remain unsuitable for reproduction for
successive years. The most vigorous populations of the fleshy-fruit
gladecress are located in areas which receive full, or near full,
sunlight at the canopy level and have limited herbaceous competition
(Hilton 1997, p. 5). Rollins (1963, p. 17) documented the loss of
fleshy-fruit gladecress individuals caused by invading grasses in an
unweeded portion of an experimental plot, while fleshy-fruit gladecress
individuals in the hand-weeded part of the plot thrived. Hilton (1997,
p. 12) was unable to relocate five populations in abandoned fields and
pastures, which McDaniel and Lyons (1987, p. 7-9) had noted as
appearing suppressed due to competition from invading weedy species.
[[Page 47129]]
Shading and competition are potential threats at the two largest
populations of fleshy-fruit gladecress (Hilton 1997, p. 68). One site,
reported to be widely open in 1968, is now partially shaded due to
closing of the canopy (Hilton 1997, p.18). Nonnative plants, including
Ligustrum vulgare (common privet) and Lonicera maackii (bush
honeysuckle), are a significant threat in many glades due to the ever
present disturbances that allow for their colonization (Hilton 1997, p.
68). Nonnative plant species pose a threat to one population of the
fleshy-fruit gladecress, where they have established near an unimproved
road traversing the site (Hilton 1997, p.18).
Under natural conditions, cedar glades are edaphically (related to
or caused by particular soil conditions) maintained through processes
of drought and erosion interacting with other processes that disrupt
encroachment of competing vegetation. Soils that develop on glades are
easily eroded, moving downslope or into fractures in the substrate. The
shallow soil, exposed rock, and frequently hot, dry summers create
xeric conditions that regulate competition and shading from encroaching
vegetation (Hilton 1997, p. 5; McDaniel and Lyons 1987, p. 6; Baskin et
al. 1986, p. 138; Rollins 1963, p. 5). Historically, periodic fires
also likely played a role in maintaining these communities (Shotz 2009,
p. 1). Extant occurrences of fleshy-fruit gladecress are primarily
located in areas modified for human use. These habitat modifications
have either eliminated or reduced the frequency of natural disturbance
processes, such as fire, that would otherwise regulate encroachment of
competing vegetation.
Residential and Industrial Development
Hilton (pers. comm., 1999) considered residential and industrial
development that had taken place in the decade prior to her study to be
the primary threat to cedar glade communities and the primary reason
for the loss of cedar glade habitat. One of the six fleshy-fruit
gladecress populations is located in the front yard of a private
residence. However, at this time, we know of no projects that would
lead to the destruction of habitat where this species is currently
located.
Climate Change
We discuss the topic of climate change in greater detail above in
the Factor A threats analysis for Short's bladderpod, which is also
applicable to the fleshy-fruit gladecress. Since, 1970, the average
annual temperature across the Southeast has increased by about 2
[deg]F, with the greatest increases occurring during the winter months.
The geographic extent of areas in the Southeast region affected by
moderate to severe spring and summer drought has increased over the
past three decades by 12 and 14 percent, respectively (Karl et al.
2009, p. 111). These trends are expected to increase. Rates of warming
are predicted to more than double in comparison to what the Southeast
has experienced since 1975, with the greatest increases projected for
summer months. Depending on the emissions scenario used for modeling
change, average temperatures are expected to increase by 4.5 [deg]F to
9 [deg]F by the 2080s (Karl et al. 2009. p. 111). While there is
considerable variability in rainfall predictions throughout the region,
increases in evaporation of moisture from soils and loss of water by
plants in response to warmer temperatures are expected to contribute to
increased frequency, intensity, and duration of drought events (Karl et
al. 2009, p. 112).
A warmer climate with more frequent droughts, but also extreme
precipitation events, may adversely affect fleshy-fruit gladecress by
altering the glade habitat the species requires. Ephemeral seeps and
streams on glades provide microhabitats important to the distribution
of the species (Hilton 1997, p. 5). Climate change may also improve
habitat conditions for invasive plant species and other plants (USFWS
2010, p. 5). A positive effect of increased drought could result from
increased mortality of woody vegetation and reduced rates of vegetation
succession.
While climate has changed in recent decades in the region where
fleshy-fruit gladecress occurs and the rate of change likely will
continue to increase for the foreseeable future, we are unable to
determine how the habitats where fleshy-fruit gladecress occurs will be
affected by these changes and how the species will respond to these
changes.
Conservation Efforts to Reduce Habitat Destruction, Modification, or
Curtailment of Its Range
The occurrence and its habitat on William B. Bankhead National
Forest (WBNF) is protected due to its location in a Native American
cultural site and the fact that cedar glade communities are considered
``rare communities'' on the WBNF and protected from detrimental effects
from agency actions (A. Cochran, U.S. Forest Service, in litt. 2005). A
thorough survey of limestone and sandstone glades on the WBNF was
completed by Schotz in 2006. Nine glades presently are known to occur
on WBNF, with sandstone glades constituting the largest percentage of
glade surface area. The fleshy-fruit gladecress inhabits Indian Tomb
Hollow Glade, the one limestone glade present on WBNF, with a surface
area of approximately 2.7 ha (1.1 ac). WBNF conducted treatment of the
nonnative invasive species Ligustrum sinense (Chinese privet) on the
Indian Tomb Hollow Glade in the fall of 2009 and summer of 2011. The
U.S. Forest Service has posted the area of the gladecress population as
closed to access and monitors impacts to the glade from off-road
vehicles. Seeds from the Indian Tomb Hollow Glade were collected in May
2010, and sent to the USDA National Center for Genetic Resources
Preservation for long-term storage.
The Service funded a survey of cedar glade habitats in the Moulton
Valley physiographic region of northwestern Alabama, the major area for
this habitat type, in the late 1990s. A survey and status update for
all fleshy-fruit gladecress populations was part of that project. The
Service recently funded surveys to update information on all
populations of this species. All sites were visited in 2006 and 2007,
and surveys continued into 2009 (Schotz 2009). This information will be
used to develop conservation measures needed to protect and enhance
populations.
Summary of Factor A
The threats to fleshy-fruit gladecress from habitat destruction and
modification are occurring throughout the entire range of the species.
These threats include agricultural conversion or incompatible
practices, maintenance of transportation rights-of-way, residential and
industrial development, and shading and competition. Conservation
efforts of the U.S. Forest Service have removed threats associated with
off-road vehicle use and encroachment of invasive species at one site;
however, maintenance of transportation right-of-ways and use of off-
road vehicles could adversely affect the remaining five extant
populations. The population-level impacts from these activities are
expected to continue into the future.
Factor B. Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
There is no information to suggest that fleshy-fruit gladecress is
collected for commercial, recreational, or educational purposes, and we
have no reason to believe that this factor will become a threat to the
species in the future.
[[Page 47130]]
Factor C. Disease or Predation
One occurrence was lost due to infection by mustard rust in the
early 1980s (Lyons and Antonovics 1991, p. 274; McDaniel and Lyons
1987, p. 11). We have no data to indicate whether this disease poses a
significant long-term threat to the species generally. There is no
information regarding predation of the species by wildlife. Grazing is
ongoing across the range of the gladecress and occurs on portions of
all extant population sites; however, there is no information to
document that cattle eat gladecress. No studies have been conducted to
investigate the effect of grazing or herbivory specifically on fleshy-
fruit gladecress.
Factor D. The Inadequacy of Existing Regulatory Mechanisms
The greatest threats to the gladecress include loss of habitat and
the plants themselves due to actions that remove the substrate under
the populations or that cover them up. These types of actions have been
associated with conversion of native glades or pastures with glades and
outcrops to other land uses and potentially herbicide applications for
the purpose of controlling invasive plants. State and Federal
regulations that might help conserve rare species on State highway
rights-of-way, including avoidance or minimization of habitat
destruction, as well as regulations that would protect plants from
herbicide applications, protect only already listed species, and
therefore do not apply to gladecress. Likewise, no existing regulations
protect the species on privately owned land, where most of the remnant
gladecress populations are found.
Factor E. Other Natural or Manmade Factors Affecting Its Continued
Existence
The fleshy-fruit gladecress is vulnerable to localized extinction
because of the small number of occurrences and the small population
sizes within the species' limited range. Small population sizes
decrease the resilience of individual fleshy-fruit gladecress
occurrences to recover from effects of other threats affecting the
species' habitat. There are only six remaining flesh-fruit gladecress
occurrences, and only one of these is protected. The loss of any
occurrences would significantly impact the species' viability by
reducing its redundancy on the landscape, which would increase its
vulnerability to stochastic environmental stressors and reduce the
species' resilience to recover from effects of threats discussed in the
above sections.
Three of the six populations of fleshy-fruit gladecress are small
in size as a result of effects of habitat loss discussed in the above
sections. The loss of populations and reductions in population sizes
have resulted in spatial isolation between these remnant populations.
These isolated populations are vulnerable to extinction by reductions
in genetic variation among the populations (Klank et al. 2012, pp. 1-2;
Shotz, pers. comm., 2013). Based on this information we conclude that
the small number of populations and the small size of populations
within the species' limited range are significant threats to fleshy-
fruit gladecress.
Cumulative Effects From Factors A Through E
Where two or more threats affect fleshy-fruit gladecress
occurrences, the effects of those threats could interact or be
compounded, producing a cumulative adverse effect that rises above the
incremental effect of either threat alone. Cumulative adverse effects
could be significant for fleshy-fruit gladecress because three of the
six extant populations are small (Factor E) and all but one of the
extant occurrences are affected by threats that result in the
destruction or modification of habitat. The vulnerability of these
occurrences to habitat modification or destruction is heightened by
effects of small population size discussed above, reduced resilience to
recover from acute demographic effects of these disturbances, and low
potential for recolonization due to isolation from other occurrences.
Proposed Determinations
We have carefully assessed the best scientific and commercial data
available regarding the past, present, and future threats to Short's
bladderpod, whorled sunflower, and fleshy-fruit gladecress. Below we
state which of the five factors are determined to be threats to these
species and summarize the severity, timing, and significance of those
threats.
Short's Bladderpod
The most significant threats to this species are described under
Listing Factors A (the present or threatened destruction, modification,
or curtailment of its habitat or range) and E (other natural or manmade
factors affecting its continued existence). Based on the Factor A
analysis, we conclude that the loss and degradation of habitat
represents the greatest threat to Short's bladderpod. Road construction
has caused the loss of habitat and all Short's bladderpod plants at
five occurrences in the past, and roadside maintenance or road widening
could adversely affect nearly 40 percent of the extant occurrences of
the species due to their position in roadside habitats. Future
development of a commuter rail project to improve intercity commute
options between the cities of Nashville and Clarksville, Tennessee,
could affect 27 percent of known extant occurrences, including some
locations where the species is found in greatest abundance.
Flooding and water level fluctuations threaten 19 percent of extant
Short's bladderpod occurrences, most notably the single Indiana
occurrence, where the species has been present in large numbers but
recently experienced a reduction in numbers due to prolonged flooding.
Overstory shading due to natural forest succession, combined with
shading and competition due to invasive, nonnative shrubs and
herbaceous species presents the most widespread, imminent threat to
Short's bladderpod, and has been implicated in the loss of several
historic occurrences. These threats are expected to continue into the
foreseeable future.
The Factor E analysis demonstrated that Short's bladderpod is
vulnerable to adverse effects of small population size, including
potential for reduced genetic variation, low numbers of compatible
mates, increased likelihood of inbreeding depression, and reduced
resilience to recover from acute demographic effects of other threats
to the species and is habitat. Fewer than 100 plants have ever been
observed at one time at 12 (46 percent) of the 26 extant occurrences,
and many of these occurrences are isolated from other occurrences.
Existing threats may be exacerbated by the effects of ongoing and
future climate change, especially projected increases in temperature
and increased frequency and severity of droughts in the Southeast and
projected increases in flooding in the Midwest.
Based on our review of the best available scientific and commercial
information, we conclude that adverse effects associated with small and
often isolated populations, as described in the Factor E analysis, both
alone and in conjunction with the widespread threats described under
Factor A, constitute significant threats to Short's bladderpod. As
discussed under Factor D, no regulatory mechanisms exist that would
prevent or restrict activities described under Factor A that constitute
significant threats to the species. Therefore, on the basis of best
available scientific and commercial information we have determined that
Short's bladderpod is in danger of extinction
[[Page 47131]]
throughout all or a significant portion of its range and that a
proposed determination as an endangered species is appropriate.
Whorled Sunflower
The most significant threats to this species are described under
Listing Factors A (the present or threatened destruction, modification,
or curtailment of its habitat or range) and E (other natural or manmade
factors affecting its continued existence). Based on the Factor A
analysis, we conclude that the loss and degradation of habitat
represents the greatest threat to whorled sunflower. Past and ongoing
risk of adverse effects from mechanical or chemical vegetation
management for industrial forestry, right-of-way maintenance, or
agriculture is a threat to three of the four extant populations of this
species. Modification of the remnant prairie habitats that the species
occupies due to shading and competition resulting from vegetation
succession also threatens these three populations, limiting growth and
reproductive output of whorled sunflower. These threats are expected to
continue in the foreseeable future. A conservation easement and
suitable habitat management currently alleviates these threats that
otherwise would adversely affect the Georgia population.
The Factor E analysis demonstrated that whorled sunflower is
vulnerable to localized extinction because of its extremely restricted
distribution and small population sizes at most known locations. Small
population size could be affecting reproductive fitness of whorled
sunflower by limiting availability of compatible mates or by causing
higher rates of inbreeding among closely related individuals. Both of
these could be contributing to reduced seed production and viability
rates, which limit the species' ability to recovery from acute
demographic effects of habitat loss or modification. The species'
dependence on remnant prairie habitats, which are isolated on the
landscape, limits the potential for recolonization in the event that
localized extinction events occur.
Based on our review of the best available scientific and commercial
information, we conclude that adverse effects associated with extremely
restricted distribution and small and isolated populations, as
described in the Factor E analysis, both alone and in conjunction with
the threats described under Factor A, constitute significant threats to
whorled sunflower. As discussed under Factor D, no regulatory
mechanisms exist that would prevent or restrict activities described
under Factor A that constitute significant threats to the species.
Therefore, on the basis of best available scientific and commercial
information we have determined that whorled sunflower is in danger of
extinction throughout all or a significant portion of its range and
that a proposed determination as an endangered species is appropriate.
Fleshy-fruit Gladecress
The most significant threats to this species are described under
Listing Factors A (the present or threatened destruction, modification,
or curtailment of its habitat or range) and E (other natural or manmade
factors affecting its continued existence). Based on the Factor A
analysis, we conclude that the loss and degradation of habitat
represents the greatest threat to fleshy-fruit gladecress. The threats
to fleshy-fruit gladecress from habitat destruction and modification
are occurring throughout the entire range of the species. These threats
include agricultural conversion for use as pasture or incompatible
practices, maintenance of transportation rights-of-way (including
mowing and herbicide treatment prior to seed set along roadsides), the
impacts of off-road vehicles, dumping, residential and industrial
development, and shading and competition. Conservation efforts of the
U.S. Forest Service have removed threats associated with off-road
vehicle use and encroachment of invasive species at one site; however,
maintenance of transportation right-of-ways and use of off-road
vehicles could adversely affect the remaining five extant populations.
Shading due to natural forest succession and competition from
nonnative invasive plants presents a significant threat to fleshy-fruit
gladecress, and has been implicated in the loss of five historic
occurrences. One site, reported to be widely open in 1968, is now
partially shaded due to closing of the canopy and the presence of
nonnative plants, including Ligustrum vulgare (common privet) and
Lonicera maackii (bush honeysuckle), and these are significant threats
in many glades due to the ever present disturbances that allow for
nonnative plant colonization. These threats are expected to continue
into the foreseeable future.
The Factor E analysis demonstrated that fleshy-fruit gladecress is
vulnerable to localized extinction because of the small number of
occurrences and the small population sizes within its limited range.
Small population sizes decrease the resilience of individual fleshy-
fruit gladecress occurrences to recover from effects of other threats
affecting its habitat and reduce genetic variation among populations.
There are only six remaining flesh-fruit gladecress occurrences, and
only one of these is protected. The loss of any occurrences would
significantly impact the species' viability by reducing its redundancy
on the landscape, which would increase its vulnerability to stochastic
environmental stressors and reduce the species' resilience to recover
from effects of threats discussed in the above sections.
Based on our review of the best available scientific and commercial
information, we conclude that adverse effects associated with limited
distribution and small population size, as described in the Factor E
analysis, both alone and in conjunction with the threats described
under Factor A, constitute significant threats to fleshy-fruit
gladecress. As discussed under Factor D, no regulatory mechanisms exist
that would prevent or restrict activities described under Factor A that
constitute significant threats to the species. Therefore, on the basis
of best available scientific and commercial information we have
determined that fleshy-fruit gladecress is in danger of extinction
throughout all or a significant portion of its range and that a
proposed determination as an endangered species is appropriate.
Significant Portion of the Range
The Act defines an endangered species as ``any species which is in
danger of extinction throughout all or a significant portion of its
range.'' A major part of the analysis of ``significant portion of the
range'' requires considering whether the threats to the species are
geographically concentrated in any way. If the threats are essentially
uniform throughout the species' range, then no portion is likely to
warrant further consideration. Based on the threats to Short's
bladderpod, whorled sunflower, and fleshy-fruit gladecress throughout
their entire known ranges, we find that these species currently are in
danger of extinction throughout all of their ranges, based on the
severity and scope of the threats described above. As discussed above,
these species are proposed for listing as endangered species, rather
than threatened species, because the threats are occurring now or will
in the near term, and their potential impacts to the species would be
severe given the limited known distribution of the species, the small
population sizes at many of the remaining sites, and the small area
occupied by many of these populations, putting these species at risk of
extinction at the present time. As these threats extend throughout
their
[[Page 47132]]
entire ranges, it is unnecessary to determine if they are in danger of
extinction throughout a significant portion of their ranges. Therefore,
on the basis of the best available scientific and commercial data, we
propose listing Short's bladderpod, whorled sunflower, and fleshy-fruit
gladecress as endangered species throughout their ranges in accordance
with sections 3(6) and 4(a)(1) of the Act.
Available Conservation Measures
Conservation measures provided to species listed as endangered or
threatened species under the Act include recognition, recovery actions,
requirements for Federal protection, and prohibitions against certain
practices. Recognition through listing results in public awareness and
conservation by Federal, State, Tribal, and local agencies; private
organizations; and individuals. The Act encourages cooperation with the
States and requires that recovery actions be carried out for all listed
species. The protection required by Federal agencies and the
prohibitions against certain activities are discussed, in part, below.
The primary purpose of the Act is the conservation of endangered
and threatened species and the ecosystems upon which they depend. The
ultimate goal of such conservation efforts is the recovery of these
listed species, so that they no longer need the protective measures of
the Act. Subsection 4(f) of the Act requires the Service to develop and
implement recovery plans for the conservation of endangered and
threatened species. The recovery planning process involves the
identification of actions that are necessary to halt or reverse the
species' decline by addressing the threats to its survival and
recovery. The goal of this process is to restore listed species to a
point where they are secure, self-sustaining, and functioning
components of their ecosystems.
Recovery planning includes the development of a recovery outline
shortly after a species is listed and preparation of a draft and final
recovery plan. The recovery outline guides the immediate implementation
of urgent recovery actions and describes the process to be used to
develop a recovery plan. Revisions of the plan may be done to address
continuing or new threats to the species, as new substantive
information becomes available. The recovery plan identifies site-
specific management actions that set a trigger for review of the five
factors that control whether a species remains endangered or may be
downlisted or delisted, and methods for monitoring recovery progress.
Recovery plans also establish a framework for agencies to coordinate
their recovery efforts and provide estimates of the cost of
implementing recovery tasks. Recovery teams (comprised of species
experts, Federal and State agencies, nongovernment organizations, and
stakeholders) are often established to develop recovery plans. When
completed, the recovery outline, draft recovery plan, and the final
recovery plan will be available on our Web site (https://www.fws.gov/endangered), or from the Service's Tennessee Ecological Services Field
Office (see FOR FURTHER INFORMATION CONTACT).
Implementation of recovery actions generally requires the
participation of a broad range of partners, including other Federal
agencies, States, Tribal, nongovernmental organizations, businesses,
and private landowners. Examples of recovery actions include habitat
restoration (e.g., restoration of native vegetation), research, captive
propagation and reintroduction, and outreach and education. The
recovery of many listed species cannot be accomplished solely on
Federal lands because their range may occur primarily or solely on non-
Federal lands. To achieve recovery of these species requires
cooperative conservation efforts on private, State, and Tribal lands.
If these species are listed, funding for recovery actions will be
available from a variety of sources, including Federal budgets, State
programs, and cost share grants for non-Federal landowners, the
academic community, and nongovernmental organizations. In addition,
pursuant to section 6 of the Act, the States of Indiana, Kentucky, and
Tennessee would be eligible for Federal funds to implement management
actions that promote the protection or recovery of Short's bladderpod.
The States of Georgia and Tennessee would eligible for Federal funds to
implement management actions that promote the protection or recovery of
whorled sunflower. Information on our grant programs that are available
to aid species recovery can be found at: https://www.fws.gov/grants.
Although Short's bladderpod, whorled sunflower, and fleshy-fruit
gladecress are only proposed for listing under the Act at this time,
please let us know if you are interested in participating in recovery
efforts for this species. Additionally, we invite you to submit any new
information on this species whenever it becomes available and any
information you may have for recovery planning purposes (see FOR
FURTHER INFORMATION CONTACT).
Section 7(a) of the Act requires Federal agencies to evaluate their
actions with respect to any species that is proposed or listed as an
endangered or threatened species and with respect to its critical
habitat, if any is designated. Regulations implementing this
interagency cooperation provision of the Act are codified at 50 CFR
part 402. Section 7(a)(4) of the Act requires Federal agencies to
confer with the Service on any action that is likely to jeopardize the
continued existence of a species proposed for listing or result in
destruction or adverse modification of proposed critical habitat. If a
species is listed subsequently, section 7(a)(2) of the Act requires
Federal agencies to ensure that activities they authorize, fund, or
carry out are not likely to jeopardize the continued existence of the
species or destroy or adversely modify its critical habitat. If a
Federal action may affect a listed species or its critical habitat, the
responsible Federal agency must enter into formal consultation with the
Service.
Federal agency actions within the species' habitat that may require
conference or consultation or both as described in the preceding
paragraph include federally funded or permitted actions occurring
within habitat for Short's bladderpod, whorled sunflower, or fleshy-
fruit gladecress (e.g., management and any other landscape altering
activities on Federal lands administered by the U.S. Army Corps of
Engineers or U.S. Forest Service; issuance of section 404 Clean Water
Act (33 U.S.C. 1251 et seq.) permits by the U.S. Army Corps of
Engineers; construction and management of gas pipeline and power line
rights-of-way by the Federal Energy Regulatory Commission; construction
and maintenance of roads or highways funded or carried out by the
Federal Highway Administration; and Federal Emergency Management
Agency-funded actions). Also subject to consultation would be provision
of Federal funds to State and private entities through Federal programs
such as the Service's Partners for Fish and Wildlife Program, State
Wildlife Grant Program, and Federal Aid in Wildlife Restoration
Program.
The Act and its implementing regulations set forth a series of
general prohibitions and exceptions that apply to endangered plants.
All prohibitions of section 9(a)(2) of the Act, implemented by 50 CFR
17.61, apply. These prohibitions, in part, make it illegal for any
person subject to the jurisdiction of the United States to import or
export, transport in interstate or foreign commerce in the course of a
[[Page 47133]]
commercial activity, sell or offer for sale in interstate or foreign
commerce, or remove and reduce the species to possession from areas
under Federal jurisdiction. In addition, for plants listed as
endangered, the Act prohibits the malicious damage or destruction on
areas under Federal jurisdiction and the removal, cutting, digging up,
or damaging or destroying of such plants in knowing violation of any
State law or regulation, including State criminal trespass law. Certain
exceptions to the prohibitions apply to agents of the Service and State
conservation agencies. The States of Georgia, Indiana, Kentucky, and
Tennessee have regulations authorizing the promulgation of lists of
endangered plants; however, with the exception of Georgia, these
regulations create no obligations on the part of landowners, public or
private, to protect State-listed plants. The Georgia Environmental
Policy Act requires that impacts to protected species be addressed for
all projects on State-owned lands, and for all projects undertaken by a
municipality or county if funded half or more by State funds, or by a
State grant of more than $250,000. The Act will, therefore, offer
additional protection to these species.
We may issue permits to carry out otherwise prohibited activities
involving endangered and threatened plant species under certain
circumstances. Regulations governing permits are codified at 50 CFR
17.62 for endangered plants, and at 17.72 for threatened plants. With
regard to endangered plants, a permit must be issued for the following
purposes: for scientific purposes or to enhance the propagation or
survival of the species.
It is our policy, as published in the Federal Register on July 1,
1994 (59 FR 34272), to identify, to the maximum extent practicable at
the time a species is listed, those activities that would or would not
constitute a violation of section 9 of the Act. The intent of this
policy is to increase public awareness of the effect of a proposed
listing on proposed and ongoing activities within the range of the
species proposed for listing. The following activities could
potentially result in a violation of section 9 of the Act; this list is
not comprehensive:
(1) Unauthorized collecting, handling, possessing, selling,
delivering, carrying, or transporting of Short's bladderpod, whorled
sunflower, or fleshy-fruit gladecress, including import or export
across State lines and international boundaries, except for properly
documented antique specimens of these taxa at least 100 years old, as
defined by section 10(h)(1) of the Act;
(2) Unauthorized removal, damage, or destruction of Short's
bladderpod or fleshy-fruit gladecress plants from populations located
on Federal land (lands owned by the U.S. Army Corps of Engineers or on
which they hold easements, or U.S. Forest Service lands); and
(3) Unauthorized removal, damage or destruction of Short's
bladderpod, whorled sunflower, or fleshy-fruit gladecress plants on
private land in violation of any State regulation, including criminal
trespass.
Questions regarding whether specific activities would constitute a
violation of section 9 of the Act should be directed to the Service's
Tennessee Ecological Services Field Office (see FOR FURTHER INFORMATION
CONTACT). Requests for copies of the regulations concerning listed
species and general inquiries regarding prohibitions and permits may be
addressed to the U.S. Fish and Wildlife Service, 105 West Park Drive,
Suite D, Athens, GA 30606; telephone 706-613-9493; facsimile 706-613-
6059.
Peer Review
In accordance with our joint policy published in the Federal
Register on July 1, 1994 (59 FR 34270), we will seek the expert
opinions of at least three appropriate and independent specialists
regarding this proposed rule. The purpose of peer review is to ensure
that our listing determination for these species is based on
scientifically sound data, assumptions, and analyses. We have invited
these peer reviewers to comment during the public comment period.
We will consider all comments and information received during the
comment period on this proposed rule during preparation of a final
rulemaking. Accordingly, the final decision may differ from this
proposal.
Public Hearings
The Act provides for one or more public hearings on this proposal,
if requested. Requests must be received within 45 days after the date
of publication of this proposal in the Federal Register. Such requests
must be sent to the address shown in the FOR FURTHER INFORMATION
CONTACT section. We will schedule public hearings on this proposal, if
any are requested, and announce the dates, times, and places of those
hearings, as well as how to obtain reasonable accommodations, in the
Federal Register and local newspapers at least 15 days before the
hearing.
Persons needing reasonable accommodations to attend and participate
in a public hearing should contact the Tennessee Ecological Services
Field Office at (931) 528-6481, as soon as possible. To allow
sufficient time to process requests, please call no later than one week
before the hearing date. Information regarding this proposed rule is
available in alternative formats upon request.
Required Determinations
Clarity of the Rule
Executive Order 12866 requires each agency to write regulations
that are easy to understand. We invite your comments on how to make
this rule easier to understand including answers to questions such as
the following:
(1) Are the requirements in the rule clearly stated?
(2) Does the rule contain technical language or jargon that
interferes with its clarity?
(3) Does the format of the rule (grouping and order of sections,
use of headings, paragraphing, etc.) aid or reduce its clarity?
(4) Would the rule be easier to understand if it were divided into
more (but shorter) sections?
(5) Is the description of the rule in the SUPPLEMENTARY INFORMATION
section of the preamble helpful in understanding the emergency rule?
What else could we do to make the rule easier to understand?
Send a copy of any comments that concern how we could make this
rule easier to understand to Office of Regulatory Affairs, Department
of the Interior, Room 7229, 1849 C Street NW., Washington, DC 20240.
You also may email the comments to this address: Exsec@ios.goi.gov.
National Environmental Policy Act (42 U.S.C. 4321 et seq.)
We have determined that environmental assessments and environmental
impact statements, as defined under the authority of the National
Environmental Policy Act of 1969, need not be prepared in connection
with listing a species as an endangered or threatened species under the
Act. We published a notice outlining our reasons for this determination
in the Federal Register on October 25, 1983 (48 FR 49244).
References Cited
A complete list of all references cited in this rule is available
on the Internet at https://www.regulations.gov under Docket No. FWS-R4-
ES-2013-0087 or upon request from the Field Supervisor, Tennessee
Ecological Services Field Office (see FOR FURTHER INFORMATION CONTACT
section).
[[Page 47134]]
Authors
The primary authors of this proposed rule are the staff members of
the Tennessee Ecological Services Field Office (see FOR FURTHER
INFORMATION CONTACT) and the Alabama Ecological Services Field Office.
List of Subjects in 50 CFR Part 17
Endangered and threatened species, Exports, Imports, Reporting and
recordkeeping requirements, Transportation.
Proposed Regulation Promulgation
Accordingly, we propose to amend part 17, subchapter B of chapter
I, title 50 of the Code of Federal Regulations, as set forth below:
PART 17--[AMENDED]
0
1. The authority citation for part 17 continues to read as follows:
Authority: 16 U.S.C. 1361-1407; 1531-1544; 4201-4245, unless
otherwise noted.
0
2. In Sec. 17.12 paragraph (h), add entries for Helianthus
verticillatus, Leavenworthia crassa, and Physaria globosa, in
alphabetical order under FLOWERING PLANTS, to the List of Endangered
and Threatened Plants, to read as follows:
Sec. 17.12 Endangered and threatened plants.
* * * * *
(h) * * *
--------------------------------------------------------------------------------------------------------------------------------------------------------
Species
-------------------------------------------------------- Historic range Family Status When listed Critical Special
Scientific name Common name habitat rules
--------------------------------------------------------------------------------------------------------------------------------------------------------
FLOWERING PLANTS
* * * * * * *
Helianthus verticillatus......... whorled sunflower... U.S.A. (AL, GA, TN) Asteraceae......... E ........... NA NA
* * * * * * *
Leavenworthia crassa............. fleshy-fruit U.S.A. (AL)........ Brassicaceae....... E ........... NA NA
gladecress.
* * * * * * *
Physaria globosa................. Short's bladderpod.. U.S.A. (IN, KY, TN) Brassicaceae....... E ........... NA NA
* * * * * * *
--------------------------------------------------------------------------------------------------------------------------------------------------------
* * * * *
Dated: July 18, 2013.
Stephen Guertin,
Acting Director, U.S. Fish and Wildlife Service.
[FR Doc. 2013-18213 Filed 8-1-13; 8:45 am]
BILLING CODE 4310-55-P